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Report of the Tomato Genetics Cooperative

Volume 63 October 2013 Report of the Tomato Genetics Cooperative Number 63‐ October 2013

University of Florida Gulf Coast Research and Education Center 14625 County Road 672 Wimauma, FL 33598 USA Foreword The Tomato Genetics Cooperative, initiated in 1951, is a group of researchers who share and interest in tomato genetics, and who have organized informally for the purpose of exchanging information, germplasm, and genetic stocks. The Report of the Tomato Genetics Cooperative is published annually and contains reports of work in progress by members, announcements and updates on linkage maps and materials available. The research reports include work on diverse topics such as new traits or mutants isolated, new cultivars or germplasm developed, interspecific transfer of traits, studies of gene function or control or tissue culture. Relevant work on other Solanaceous species is encouraged as well.

Paid memberships currently stand at approximately 50 from 16 countries. Requests for membership (per year) at US$10 for the online edition should be sent to Dr. J.W. Scott, [email protected]. Please send only checks or money orders. Make checks payable to the University of Florida. We are sorry but we are NOT able to accept cash, wire transfers or credit cards.

Cover: This is a figure from the 2013 feature article entitled “Using fluorescence in situ hybridization (FISH) to improve the assembly of the tomato genome” by Shearer et al. describing work that involved six laboratories at several institutions. A more detailed refereed paper will be published later, but you can get a glimpse of this exciting work right here in the Tomato Genetics Cooperative so be sure to check it out.

Page 1 of 64 TABLE OF CONTENTS TGC REPORT VOLUME 63, 2013

Foreword ………………………………………………………………………...... ………..1 Announcements ………………………………………………………………………………………………………… 3 Feature Article Using fluorescence in situ hybridization (FISH) to improve the assembly of the tomato genome Lindsay A. Shearer, Lorinda K. Anderson, Hans. de Jong, Sandra Smit, José Luis Goicoechea, Bruce A. Roe, Axin Hua, James J. Giovannoni, and Stephen M. Stack ……………………………………………………………………………………………………………………..……5 Research Reports Evaluation of Near Isogenic Tomato Lines with and without the Bacterial Wilt Resistance Allele, Bwr-12 Peter Hanson, Chee-Wee Tan, Fang-I Ho, Shu-Fen Lu, Dolores Ledesma and Jaw-Fen Wang ………………………………………………………………………………………………………..………15 Distribution of major QTLs associated with resistance to Ralstonia solanacearum phylotype I strain in a global set of resistant tomato accessions Fang-I Ho, Chi-Yun Chung, and Jaw-Fen Wang ………………………………………….…….…22 Varietal Pedigrees Fla. 8111B; a large-fruited, globe shaped tomato breeding line J.W. Scott ……………………………………………………………………………………………………….…..31 Fla. 8233; a germplasm line with partial resistance to bacterial spot races T1, T2, T3, T4 and Xanthomonas gardneri. J.W. Scott and S.F. Hutton …………………………………………………………………………………. 32 Stock Lists Revised List of Wild Species Stocks Chetelat, R. T.………………………………………………………...... 33 Membership List ……………………………………………………………………………………………….…….. 61 Author List …………………………………………………………………………………..………………….….…... 64

Page 2 of 64 ANNOUNCEMENTS TGC REPORT VOLUME 63, 2013

From the editor:

Happy Halloween 2013 to all TGC members! Belated Happy Halloween to all you non-members. Volume 63 will be online in October this year which makes two TGC volumes in one year since the 2012 TGC did not get published until March 2013. Pretty spooky!! Thanks to those who sent in papers this year. This year’s feature article by Steve Stack et al., on the use of FISH to improve the tomato genome assembly will be published in greater detail later in a high impact journal, but you got a sneak peek first right here at TGC. Perhaps this will stimulate some of you to do the same type of thing in future years.

Big News on the keyword search function: All Volumes except the most recent one each year of the Tomato Genetics Cooperative are now all searchable by keyword and this will allow you to efficiently find articles about topics of interest. We have been striving for this for years and it is finally a reality! Follow instructions at the top of the Online Volumes section on the TGC website. When you get the list of articles the ones with the .html (or .htm) extension will be individual articles. Those with .pdf extensions will pull up the whole volume. The browser will give you the volume so citations can be easily tracked down. If anyone has any suggestions for improvement of this feature or anything else about TGC please contact me. Thanks to Dolly Cummings for her expert help with TGC operations and to Christine Cooley for her help with the website. Dolly and Christine are also responsible for getting the keyword search functioning. The value of great employees cannot be overstated!

I hope you have a good year ahead, maybe the US government will be open again by next year, but given the present political scene that is not something to be counted on (it may open again but I’m sure Congress will find a way to shut it down again). Now that is spooky!

My contact information:

Jay W. Scott, Ph.D. Gulf Coast Research & Education Center 14625 CR 672 Wimauma, FL 33598 USA Phone; 813-633-4135 Fax: 813-634-0001 Email: [email protected]

Jay W. Scott Managing Editor

Page 3 of 64 ANNOUNCEMENTS TGC REPORT VOLUME 63, 2013

Upcoming Meetings

XVIIIth EUCARPIA Meeting of the Tomato Working Group, April 22- 25, 2014 https://colloque.inra.fr/eucarpia2014-tomato-avignon

Tomato Breeders' Round Table 14 - 17, 2014 Asheville, NC. Check Tomato Genetics Cooperative website http://tgc.ifas.ufl.edu/index.htm for further details.

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Using fluorescence in situ hybridization (FISH) to improve the assembly of the tomato genome

Lindsay A. Shearer1, Lorinda K. Anderson1, Hans. de Jong2, Sandra Smit3, José Luis Goicoechea4, Bruce .A Roe5, Axin Hua5, James J. Giovannoni6, and Stephen M. Stack1

1Department of Biology, Colorado State University, Fort Collins, CO 80525, USA 2Laboratory of Genetics, Wageningen University, Radix‐West, W1.Cc.054, Droevendaalsesteeg 1,6708 PB Wageningen, The Netherlands 3Laboratory of Bioinformatics, Wageningen University, Droevendaalsesteeg1, 6708 PB Wageningen, The Netherlands 4Arizona Genomics Institute, University of Arizona, Tucson, AZ 85721 5Department of Chemistry and Biochemistry, Stephenson Research and Technology Center, University of Oklahoma, Norman, OK, USA73019 6Department of Plant Biology, Cornell University, Ithaca, NY 14853, USA

INTRODUCTION The genome of tomato (Solanum lycopersicum, 2n = 2x = 24, cultivar Heinz 1706) recently was described in Nature (The Tomato Genome Consortium 2012) as twelve DNA pseudomolecules that correspond to the twelve tomato chromosomes. Ideally the DNA sequence of a pseudomolecule is the same DNA sequence that runs from the head (end of the short arm) to the tail (end of the long arm) of the chromosome. In reality, most pseudomolecules are not complete but consist of a series of more or less long DNA sequences called scaffolds that are assemblies obtained from matching overlapped segments of sequenced DNA. A pseudomolecule is assembled by lining up its scaffolds in the same order and orientation they would have in the corresponding chromosome. Two methods were used to order and orient (arrange) DNA scaffolds in tomato pseudomolecules. One relies on the tomato Kazusa EXPEN 2000 linkage map (Shirasawa et al. 2010; http://solgenomics.net/cview/map.pl?map_version_id=103), which is a high‐resolution linkage map based on an F2 mapping population derived from an interspecific cross between the Solanum lycopersicum cultivar LA925 and the wild species S. pennellii line LA716 (Fulton et al. 2002). Mapped DNA sequences were identified in scaffolds, and then scaffolds were ordered and oriented according to the locations of these sequences in the linkage map. This method should be relatively accurate in parts of the genome where crossing over is frequent, i.e., distal euchromatin, but it is expected to be less accurate in regions of the genome where crossing over is infrequent, e.g., pericentric heterochromatin. The other method uses fluorescence in situ hybridization (FISH) of bacterial artificial chromosomes (BACs) to locate mostly single copy DNA sequences on spreads of synaptonemal complexes (SCs = pachytene chromosomes; Figure 1). Each BAC contains an insert of about 100 kb of tomato genomic DNA. While the two methods usually yield the same arrangement of scaffolds in distal euchromatin, they often do not agree in pericentric heterochromatin that includes 77% of the tomato genome (Peterson et al. 1996) and about 10% of tomato’s ~35,000 nuclear genes (Van der Hoeven et al. 2002; Wang et al. 2006; The Tomato Genome Consortium 2012; Peters et al. 2009). In cases of disagreement, the published paper on the tomato genome described scaffold

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order and orientation according to the linkage map (The Tomato Genome Consortium 2012). At the time of publication, this was appropriate because most of the needed FISH localizations at scaffold borders were not yet available. However, these FISH localizations have now been completed for all tomato scaffolds, and numerous differences were noted between the linkage map‐based scaffold arrangements compared to the FISH‐based scaffold arrangements. We present evidence that FISH results are preferable for arranging scaffolds, especially in portions of the genome where the linkage map is inaccurate due to low rates of crossing over, e.g., pericentric heterochromatin.

MATERIALS AND METHODS Two lines of tomatoes Solanum lycopersicum, var. Cherry, accession LA4444 and Solanum lycopersicum, var. Heinz 1706 were grown from seeds to flowering in a greenhouse with controlled temperature and supplemental lighting. While the genome of tomato variety Heinz 1706 was sequenced (The Tomato Genome Consortium 2012), we used tomato variety Cherry, accession LA4444 for spreading SCs because it flowers abundantly and has indeterminate growth, unlike Heinz 1706. Based on FISH, the two genomes seem to be structurally almost identical. Tomato SC spreads. Spreads were prepared for FISH as described (Stack et al. 2009; Stack and Anderson 2009). Cot 100 nuclear DNA was used for chromosomal in situ suppression (CISS) hybridization because it includes most of the repetitive sequences in the tomato genome (Peterson et al. 1997; Zwick et al. 1997; Chang 2004; Peterson et al. 1998). Probes for fluorescence in situ hybridization (FISH) were prepared from tomato HindIII, MboI, EcoRI, and sheared BAC libraries and from a tomato fosmid library all located at Cornell University (http://www.sgn.cornell.edu). The DNAs were labeled with digoxygenin, biotin, or dinitrophenol (DNP) using a nick translation kit (Roche Applied Science). FISH. FISH was performed as described (Zhong et al. 1996; Chang 2004). Briefly, SC spreads on glass slides were fixed briefly in 45% acetic acid followed by 1:3 acetic ethanol and then digested with RNase followed by pepsin. After additional fixation in 1% paraformaldehyde, labeled probes with Cot 100 tomato DNA in standard hybridization medium was placed on each slide. Slides were incubated at 80° to denature the DNA and then incubated at 37° to hybridize the DNA. Slides were washed in conditions for 80% stringency, i.e., hybridization of DNAs with at least 80% complementarity (Schwarzacher et al. 2003). After blocking, slides were incubated with appropriate antibodies for the probes (mouse anti‐biotin, biotinylated donkey anti‐mouse, rat anti‐DNP, and/or sheep anti‐digoxigenin) followed by incubation with appropriate secondary antibodies labeled with fluorescent dyes (TRITC, Dylight 649, and FITC). After immunolabeling, cover glasses were mounted with Vectashield (Vector laboratories) containing 5 µg/ml 4’,6‐diamidino‐2‐phenylindole (DAPI). Microscopy and photography were performed with Leica DM 5000B and DM 5500B microscopes, both equipped for phase contrast and fluorescence microscopy with DAPI, FITC, TRITC, and Cy5 filter cubes and zero pixel shift. Images were captured with cooled Hamamatsu monochrome 1344X1044 pixel cameras using IP Lab software. Measuring positions of BACs on SCs. See Chang et al. (2007) and Stack et al. (2009) for details. The position of a BAC was measured on ten or more different SC spreads (with a few rare exceptions) to yield an average position expressed as a percent of the arm length from the

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kinetochore. This decimal fraction was multiplied by the average length of the arm in micrometers to describe the position of the BAC in micrometers from the kinetochore (Chang et al. 2007; http://solgenomics.net/cview/map.pl?map_version_id=25). Optical mapping. Optical mapping of tomato nuclear DNA was performed by OpGen, Inc. using the protocol previously described for other plant genomes (Zhou et al. 2009; Dong et al. 2013; Young et al. 2011). Optically mapped scaffolds were aligned with sequenced scaffolds by comparing patterns of digestion by BamH I of the former with expected in silico cutting patterns of the latter using OpGen‐supplied Genome‐Builder™ software. When optical mapping spanned gaps between sequenced scaffolds, superscaffolds were created and the size of their included gaps were estimated.

RESULTS AND DISCUSSION For most BACs, FISH on SC spreads resulted in a single, discrete site of hybridization (Figure 1). Because there is little or no distortion of SCs in spreads, BAC positions can be mapped with a high degree of accuracy. In addition, with inclusion of Cot 100 DNA for CISS hybridization to block repeated sequences, BAC localizations in heterochromatin often were as informative as those in euchromatin. We have localized 627 BACs to unique sites on tomato SCs, including sites in euchromatin, heterochromatin, chromomeres, and kinetochores (see http://solgenomics.net/cview/map.pl?map_version_id=25 for localization and identification of most of these BACs on a pachytene idiogram with supporting FISH images.

Figure 1. Example of a complete set of spread tomato SCs used for FISH. A. Digitally reversed phase contrast image of a complete set of 12 SCs before FISH. SCs are thicker in distal euchromatin and thinner in proximal pericentric heterochromatin. Kinetochores are white ellipsoids about 1 micrometer in diameter near the center of pericentromeric heterochromatin of each SC. The kinetochore of SC 7 is marked “7.” B. Fluorescent image of the same SC set shown in A after DAPI staining and FISH. Pachytene chromosomes are fuzzy blue structures due to chromatin loops extending laterally from the SC axes. Kinetochores are visible as intense blue ellipsoids, one of which is marked “7” on chromosome 7. Note the red foci in the distal euchromatin of the long arm of chromosome 7, which mark the location of BAC LE_HBa0227C07. C. Enlarged digitally reversed phase image of the same SC 7 shown in A and B. The borders of distal euchromatin and pericentric heterochromatin in both arms are marked with thin transverse white lines. Red fluorescent foci in the distal euchromatin of the long arm of chromosome 7 mark the location of BAC LE_HBa0227C07. The bar in A represents 10 µm for A and B. The bar in C represents 2 µm.

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Comparing linkage map‐based and FISH‐based pseudomolecules. Scaffolds in each pseudomolecule originally were numbered (1, 2, 3, …) based on the order determined from the Kazusa EXPEN 2000 linkage map starting from the head (= end of the short arm) of the chromosome (The Tomato Genome Consortium 2012). To determine the order and orientation of scaffolds by FISH, BACs at or near each end of most scaffolds were localized by FISH to SC spreads, although only one BAC was localized for very short scaffolds. This allowed us to determine the order and orientation of scaffolds independently from the linkage map and to estimate gap sizes between scaffolds. For example, Figure 2 shows FISH localization of four BACs that mark each end of two adjacent scaffolds on the short arm of chromosome 3. Because scaffold numbering is based on the linkage map, the two adjacent scaffolds are numbered 1 and 4. Scaffolds 2 and 3 were localized by FISH to separated positions on the long arm of chromosome 3. The SC lengths corresponding to each scaffold (e.g., the distance between the purple and green signals for scaffold 1 and between the red and turquoise signals for scaffold 4) and the SC lengths corresponding to the gaps between scaffolds (e.g., the distance between the green and red signals for gap 1‐4) were measured for all 91 scaffolds and all 79 gaps distributed along the twelve tomato SCs.

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Figure 2. Reversed phase image of the short arm of SC3 overlaid with fluorescent signals from four BACs that were localized simultaneously by FISH. The four BACs define the edges of two scaffolds and the gap between them. The upper lobe of the white, dumbbell‐shaped structure to the right is the kinetochore, while the lower lobe is debris (visible by phase microscopy but not by fluorescence). BAC SL_s0009C01 (purple) is at the head (H = toward the end of the short arm) and BAC SL_s0086D22 (green) is at the tail (T = toward the end of the long arm) of scaffold 1 (= SL2.40sc04439). BAC SL_s0018K15 (red) is at the head and BAC SL_s0002G24 (turquoise) is at the tail of the adjacent scaffold 4 (= SL2.40sc4696). Scaffolds 1 and 4 are in distal euchromatin. The space between the green signals and the red signals is the gap between scaffolds 1 and 4. The purple and turquoise foci mark the location of DNP‐labeled BAC probes that were the same color in the original image but which have been given different colors here. In the diagram the thick grey segments labeled 1, 4, 5, and 6 represent scaffolds SL2.40sc04439, SL2.40sc4696, SL2.40sc05330, and SL2.40sc4126, respectively; with their lengths proportional to the amounts of DNA they represent. BAC‐FISH localizations used to order and orient scaffolds 5 and 6 are not illustrated. Scaffold numbering is based on the order of scaffolds determined from the Kazusa EXPEN 2000 linkage map. Based on FISH, these scaffolds have the same head‐tail orientation, but a different order from that derived from the linkage map (Figures 3, 4). Gaps between scaffolds are named according to the scaffolds on either side, e.g., 1‐4, 4‐5, etc, and their lengths (white lines between the scaffolds) are proportional to the amount of DNA they are estimated to represent based on measurements of the relative positions of the four foci on 10 or more SCs. The upper bar represents 1 µm in reference to the SC segment, while the lower bar represents 2 Mb in reference to the pseudomolecule.

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For 46 of the scaffolds, FISH‐based arrangements were the same as linkage map‐based arrangements (Figure 3; thick black bars). However, for the remaining 45 scaffolds representing 34% (259.8 Mb) of the sequenced DNA (760.0 Mb), the FISH‐based arrangement differs from the linkage‐based arrangement (Figure 2; thick colored bars). Twenty‐eight of the differences involved only order (red bars), three differences involved only orientation (blue bars), and fourteen differences involved both order and orientation (purple bars). In general, scaffolds located in euchromatic regions of the chromosomes were more likely to be in the same arrangement in both linkage map‐based and FISH‐based pseudomolecules, while scaffolds located in pericentric heterochromatin were more likely to differ.

Figure 3. Diagrammatic representations of the twelve tomato pseudomolecules containing 91 scaffolds showing the order and orientation of scaffolds based on FISH compared to the linkage map. Thick segments represent sequenced scaffolds, and thin black lines connecting adjacent scaffolds represent gaps. All scaffolds and gaps are proportional in length to the amount of DNA they represent. The 46 scaffolds shown in dark gray have the same order and orientation using both FISH and linkage methods, while the 45 colored scaffolds have different orders and/or orientations. The dark gray scaffolds represent 66% (500.2 Mb) of the sequenced genome and strongly tend to be distal and involve euchromatin. The colored scaffolds represent 34% (259.8 Mb) of the sequenced genome and tend to be proximal and involve heterochromatin. The 28 red scaffolds were changed in order only, the three blue scaffolds were changed in orientation only, and the 14 purple scaffolds were changed in both order and orientation. The bar represents 10 Mb.

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Figure 4 shows a more detailed comparison of the linkage map‐based pseudomolecule and the FISH‐based pseudomolecule for chromosome 3. Of the 13 scaffolds, nine (2‐9, 12) differ in order and/or orientation. The most notable discrepancies include scaffold 2 that was placed in the euchromatin of the short arm by linkage mapping but was located by FISH in the heterochromatin of the long arm, scaffold 3 (a small scaffold of only ~ 400 kb) that was placed in the short arm near/in the heterochromatin but located by FISH in the euchromatin of the long arm, and scaffold 12 that was located in the middle of the euchromatic portion of the long arm by linkage mapping but located by FISH in the heterochromatin of the long arm. In addition, the kinetochore moves from scaffold 6 in the map‐based pseudomolecule to the gap between scaffolds 6 and 9 in the FISH‐based pseudomolecule.

Figure 4. Diagrammatic representations of tomato SC3 (A) with corresponding linkage map‐ based (B) and FISH‐based (C) pseudomolecules. (A) The SC is represented as a horizontal black line with a black ellipsoid indicating the position of the kinetochore. The SC is oriented with its short arm at the left. Pericentric heterochromation is represented as a grey layer to either side of kinetochore, and the approximate location of this heterochromatin is projected onto the DNA pseudomolecules below. In the pairs of pseudomolecules (B and C), thick segments represent sequenced scaffolds, and gaps between scaffolds are represented by thin black lines. Arrowheads in the scaffolds indicate the orientation of each scaffold with regard to the linkage map from head to tail (i.e. toward the end of the long arm of the chromosome), and numbers above scaffolds indicate their order from the head to the tail of the pseudomolecule based on the linkage map. Because the linkage map was used to define scaffold order and orientation, the scaffolds in the linkage map‐based pseudomolecule (B) are always in the “correct” order and orientation while the corresponding FISH‐based pseudomolecule (C) differs in several respects from the linkage‐based pseudomolecule. Scaffolds that are in the same order and orientation in the two pseudomolecules are black, while scaffolds that are different are presented in matching colors (along with scaffold numbers). Changes in order (position) are obvious and changes in orientation are shown by red arrowheads that are reversed in direction in the FISH‐based pseudomolecules. In linkage‐based pseudomolecules, all gaps are shown the same width because the linkage map is not useful for estimating gap sizes. In FISH‐based pseudomolecules, the lengths of gaps are proportional to the amount of DNA estimated to be in the gaps. Names for scaffolds 1, 4, 5, and 6 are given in Fig. 1. Scaffolds 2, 3, 7 – 13 are named, respectively, SL2.40sc04822, SL2.40sc06911, SL2.40sc06725, SL2.40sc04704, SL2.40sc04616, SL2.40sc03806, SL2.40sc03796, SL2.40sc03721, and SL2.40sc03701. The upper bar represents 1 micrometer of SC length (A), and the lower bar represents 5 megabases for each pair of pseudomolecules (B and C).

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Optical mapping strongly supports FISH‐based scaffold arrangement. Optical mapping is an independent physical method for ordering and orienting scaffolds, which has been used for genome assembly in other higher eukaryotes (Young et al. 2011; Zhou et al. 2009; Dong et al. 2013). Using this method, we were able to order and orient 38 of the 91 tomato scaffolds. Optical mapping agreed with FISH results for all of these 38 scaffolds (100%), but agreed with map‐based results for only 22 of these (58%). FISH is useful for determining the sizes of gaps between scaffolds in pseudomolecules. The amounts of DNA in gaps between adjacent scaffolds in pseudomolecules were estimated by multiplying the gap lengths in micrometers of SC by the linear density of DNA appropriate for the chromatin type (euchromatin = 1.5 Mb/µm, heterochromatin = 8.1Mb/µm, kinetochores = 3.3 Mb/µm). The total amount of DNA estimated to be in gaps is 38.5 Mb or (38.5 MB/917 Mb =) roughly 4% of the genome (Michaelson et al. 1991). Individual gaps range in size from a high of 2.9 Mb to gaps too small to estimate by FISH. Chromosome 0 BACs localize by FISH to gaps between scaffolds. Chromosome 0 BACs are defined as BACs that have been fingerprinted, partially sequenced, and/or completely sequenced but which do not fit into any of the twelve pseudomolecules because they lack both alignment with scaffold sequences and have no mapped marker sequences. Of the 93 chromosome 0 BACs that we have FISHed, 75 were successfully localized. Remarkably, 93% (70) of these were located in gaps between scaffolds.

CONCLUSIONS FISH has improved the assembly of the tomato genome by correcting the order and/or orientation of sequenced scaffolds that originally were aligned using linkage maps. Most, but not all, of these changes were observed in heterochromatic regions, where linkage maps are inaccurate due to the low rate of recombination. Optical mapping, an independent physical method to assess genome sequence assembly, was useful in addressing the arrangement of 38 of the 91 scaffolds, and those results invariably agreed with FISH results. We estimated DNA density in different parts of the genome (euchromatin, heterochromatin and kinetochores) and, from that, calculated the amount of DNA in gap between scaffold sequences. In total, we estimate that only about 4% of the tomato genome remains un‐sequenced. Chromosome 0 BACs that did not fit into any of the sequenced scaffolds were localized by FISH almost exclusively to gaps between scaffolds. These BACs are seed BACs to help complete the genome sequence. Finally, FISH‐based order and orientation of scaffolds should be used to improve the linkage map for tomato.

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ACKNOWLEDGEMENTS This work was supported by National Science Foundation Grants DBI‐0116076, DBI‐0421634, and DBI‐0820612 (J.J.G., B.A.R., S.M.S.); the Centre for BioSystems Genomics (CBSG) which is part of the Netherlands Genomics Initiative/Netherlands Organization for Scientific Research and by the European Commission (EU‐SOL project PL 016214) (H. de J.); European Union FP6 Integrated Project EU‐SOL PL 016214 (S.S.). This paper is an abbreviated version of a larger manuscript in preparation.

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Page 14 of 64 RESEARCH REPORTS TGC REPORT VOLUME 63, 2013

Evaluation of Near Isogenic Tomato Lines with and without the Bacterial Wilt Resistance Allele, Bwr-12

Peter Hanson, Chee-Wee Tan, Fang-I Ho, Shu-Fen Lu, Dolores Ledesma and Jaw-Fen Wang

AVRDC – The World Vegetable Center, 60 Yi-Min Liao, Shanhua, Tainan, Taiwan

Introduction Bacterial wilt (BW) of tomato, caused by Ralstonia solanacearum, is one of the most important diseases limiting tomato production in the tropics (Hayward, 1991; Wang et al., 1998). The broad host range and vast genetic diversity of the pathogen as well as its widespread distribution complicate disease control. Resistant cultivars would provide farmers a cheap and effective means of control. However, the development of resistant cultivars has been difficult because resistance is often multigenic (Danesh et al., 1994; Carmeille et al., 2006; Wang et al., 2013), and its expression is often incomplete and strongly affected by pathogen strain and other environmental factors such temperature, soil moisture and pH. Resistance has been associated with negative horticultural traits such as small fruit size (Hayward, 1991; Hanson et al., 1996; Scott et al., 2005).

Effective screening and selection for BW resistance has challenged tomato breeders for many years. Screening segregating populations in pathogen-infested fields (sick plots) is often unreliable because of uneven pathogen distribution and differences in environment. AVRDC and some other programs use greenhouse seedling screening techniques to manage some sources of environmental variation, but differences in microenvironment in the greenhouse may cause variable responses. Furthermore, BW screening is most effective when mean temperatures are ≥ 25 °C, which limits screening to certain periods of the year unless heated greenhouses are available. Molecular markers linked to BW resistance genes would improve the effectiveness of BW screening, and help identify and combine key BW resistance genes in the same cultivar. Wang et al. (2013) identified two major quantitative trait loci (QTLs) in tomato cultivar ‘Hawaii 7996’ (H7996) associated with stable BW resistance: Bwr-12 on chromosome 12 and Bwr-6 on chromosome 6. Bwr-12, delimited by simple sequence repeat (SSR) markers SLM12-2 and SLM12-10, is important for resistance to Phylotype 1 (Asia) pathogen strains (Wang et al. 2013; Geethanjali et al., 2011). The objective of this study was to assess the gain in resistance due to Bwr-12 by comparing near isogenic lines with and without Bwr-12 and to confirm the usefulness of SLM12-2 and SLM12-10 for marker-assisted selection.

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Materials and Methods

Plant materials. BW-resistant AVRDC line CLN2585D was crossed to BW-susceptible parent G2 in 2007 with the cross code CLN3125. BW resistance in CLN2585D was not derived from H7996 and originated from several possible sources, including lines from the University of the -Los Baños, cultivars ‘Venus’ and ‘Saturn’ from North Carolina State University, CRA-coded lines bred by the French National Institute for Agricultural Research (INRA) in Guadeloupe, French West Indies, and others (Daunay et al., 2010). Generation advance of CLN3125 from the F2 to F7 took place at AVRDC Taiwan from 2007-2010 following pedigree selection. During generation advance, segregating populations and lines of CLN3125 were screened and selected for Tomato yellow leaf curl disease resistance, fruit set, and fruit firmness, color, shape and size. BW reactions of CLN3125-coded lines were assessed in the greenhouse by drench inoculation (described below) in the F5 (May-June 2009) and F7 generations (May-June 2010). After SSR markers SLM12-2 and SLM12-10 became available, parents CLN2585D and G2, 6 F7-derived F8 lines (F7:8) developed from CLN3125 were tested for presence of Bwr-12 along with resistant check H7996. Marker analysis revealed that lines CLN3125A-23 and CLN3125L were heterogeneous (mixture of Bwr-12 homozygotes and heterozygotes). CLN3125A-23 and CLN3125L originated from the F2 plant but their pedigrees diverged at the F3.

Development of Bwr-12 near-isogenic lines. From January-March 2011, 108 and 122 F8 seedlings, respectively, from CLN3125A-23 and CLN3125L were grown in a plastic house and individually genotyped for Bwr-12 with SLM12-2 and SLM12-10. Primer sequences were: PCR Marker Forward primer Reverse primer size (bp) SLM12-2 ATCTCATTCAACGCACACCA AACGGTGGAAACTATTGAAAGG 209 SLM12-10 ACCGCCCTAGCCATAAAGAC TGCGTCGAAAATAGTTGCAT 242

Based on marker results, plants homozygous for Bwr-12 or the susceptible allele were identified, transplanted to the field, and harvested individually to create near-isogenic lines (NILs). Within CLN3125A-23, 22 resistant and 19 susceptible NILs were developed; within CLN3125L, 23 resistant and 21 susceptible NILs were produced. NILs derived from CLN3125A- 23 and CLN3125L were assigned to NIL Groups 1 and 2, respectively.

NIL evaluation for bacterial wilt reaction. Inoculations were conducted with virulent Ralstonia solanacearum strain Pss4, a Phylotype 1 (Asia), race 1 isolate from Taiwan (Wang et al., 2013). Pss4 has been used in routine AVRDC BW screening for many years. Four-week-old seedlings, approximately in the five-leaf stage, were inoculated by pouring 20 ml of 1 x 108 inoculum on the soil surface at the base of each plant. Plant roots were not wounded before

Page 16 of 64 RESEARCH REPORTS TGC REPORT VOLUME 63, 2013 inoculation. The number of wilted plants in each plot was counted weekly for four weeks beginning one week after inoculation and the percentage of healthy plants per plot was determined after the last evaluation. Plots included 20 plants and entries were replicated twice and arranged according to a randomized complete block design. Entries within NIL Groups 1 and 2 were randomized separately but screened in the same greenhouse at the same time. Checks included in both NIL groups were parents CLN2585D and G2, CLN3125A-23 and CLN3125L, H7996 (homozygous for Bwr-12 and Bwr-6) and L390 (AVRDC susceptible check). Data were subjected to analysis of variance over years using the General Linear Models procedure of Statistical Analysis Systems software. Percent survival means for each plot were transformed by arc sine before analysis. Contrasts were constructed to compare group means of resistant and susceptible NILs within NIL groups.

Results and Discussion

The combined analysis of variance over years revealed highly significant means squares for entry and entry-by-year interactions for each NIL Group (Table 1). The entry-by-year interaction arose mainly from changes in the magnitude of differences among entries between years and not from major rank changes. Contrasts between group means of resistant (homozygous for Bwr-12) versus susceptible (homozygous for susceptible allele) NILs were highly significant within NIL groups. However, contrasts between group means of resistant NILs versus CLN3125A-23, and susceptible NILs versus CLN3125A-23 in NIL Group 1, and the susceptible NILs versus CLN3125L in NIL Group 2 were non-significant. Only the contrast of the group mean of resistant NILs versus CLN3125L in NIL Group 2 was significant. Variations within resistant NILs or susceptible NILs were not statistically significant.

Bacterial wilt pressure in the greenhouse trials was high, evidenced by the low percent survival means of the susceptible check L390, susceptible parent G2, and also the relatively high percentages of wilted plants in resistant check H7996 (14%) and resistant parent CLN2585D (~30%) (Table 2). The difference in percent survival between NILs with or without Bwr-12 was 26.1% and 35.7% in NIL Groups 1 and 2, respectively. These differences are slightly lower than the 45% difference (reported as mean percentage of wilted plants) between recombinant inbred line (RIL) groups with or without Bwr-12 and tested with Taiwan strain Pss4b (Wang et al., 2013). In this study, the ranges in mean percent survival of resistant NILs were wide (Table 2) but did not overlap with the ranges of the susceptible NIL means. It is interesting that the mean percent survival of CLN2585D was significantly greater than almost all resistant NILs; only one NIL in NIL Group 2 was not significantly different in percent survival from CLN2585D according to the Waller-Duncan k-ratio t test (data not shown).

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Although CLN2585D does not possess Bwr-6, it is possible that it carries one or more minor BW resistance genes not present in CLN3125A-23 and CLN3125L.

It would seem that CLN2585D carries Bwr-12 even though H7996 was not among its progenitors. Given the extensive exchange of BW resistant lines and sources among tomato breeding programs over many years (Daunay et al., 2010), it is not too surprising that Bwr-12 was introduced and incorporated into some AVRDC lines. SSR markers SLM12-2 and SLM12-

10 flanking Bwr-12 are effective and inexpensive, enabling BW selection to occur in the F2 generation. Marker-assisted selection for Bwr-12 and other BW resistance genes has facilitated early elimination of susceptible plants, reduced the number of BW greenhouse confirmation screening trials during generation advance, and has enabled characterization of lines for the presence of specific resistance genes.

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Table 1. Combined analysis of variance1 over years for NILs with or without Bwr-12 and checks screened for bacterial wilt reaction in the greenhouse, AVRDC, May-June 2011 and May-June 2013. NIL Group 1 DF Mean square F Value Year 1 1722.69 Reps (Year) 2 408.21 Entry 46 1001.53 6.39** Resistant NILs 21 158.69 1.01ns Susceptible NILs 18 117.57 0.75ns Resistant vs Susceptible NILs 1 24924.52 159.13** Resistant 1 156.21 1.00ns NILs vs. CLN3125A-23 Susceptible 1 187.48 1.20ns NILs vs. CLN3125A-23 Others 4 3307.60 21.12** Entry*Year 46 156.63 1.78** Error 92 88.08

NIL Group 2 DF Mean square F Value Year 1 4.38 Reps (Year) 2 207.40 Entry 49 1221.21 8.31** Resistant NILs 22 160.92 1.09ns Susceptible NILs 20 77.65 0.53ns Resistant vs Susceptible NILs 1 37148.17 252.76** Resistant NILs vs. CLN3125L 1 291.49 1.98* Susceptible NILs vs. CLN3125L 1 0.49 0.00ns Others 4 3911.21 26.61** Entry*Year 49 146.97 1.59* Error 98 92.66 1Percent survival data were transformed by arcsine before analysis. *,** Significant at P<0.05 and P< 0.01, respectively. ns non-significant

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Table 2. Percent survival group means of near-isogenic lines with or without Bwr-12, and checks assessed for bacterial wilt reaction in the greenhouse, AVRDC, Taiwan NIL NIL Group/check SLM SLM Entry Range Mean Group 1 12-2 12-10 no. (% survival)1 NILs-Resistant ++ ++ 22 18.7–47.3 30.2 NILs- Susceptible -- -- 19 0.0–13.9 4.1 Difference 26.1**

H7996 ++ ++ 1 86.1 a CLN2585D ++ ++ 1 64.8 a CLN3125A-23 H H 1 21.1 bc L390 -- -- 1 6.4 cd G2 -- -- 1 0.0 d

NIL NILs-Resistant ++ ++ 23 19.8–64.5 42.4 Group 2 NILs- Susceptible -- -- 19 1.3–11.8 6.7 Difference 35.7**

H7996 ++ ++ 1 86.3 a CLN2585D ++ ++ 1 74.8 a CLN3125L H H 1 5.2 b L390 -- -- 1 1.3 b G2 -- -- 1 0.0 b

1Mean percent survival four weeks after drench inoculation with R. solanacearum Pss4 2++, = homozygous for Bwr-12 allele; -- = homozygous for the susceptible allele; H= heterogeneous *,** Significant at P<0.05 and P< 0.01, respectively. ns non-significant Means within columns and NIL groups followed by the same letter are not significantly different by Waller-Duncan

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References: Carmeille, A., C. Caranta, J. Dintinger, P. Prior, J. Luisetti and P. Busse. 2006. Identification of QTL for Ralstonia solanacearum race 3-phylotype II resistance in tomato. Theor Appl Genet 113: 110–121.

Danesh D., S. Aarons, G.E. McGill and N.D. Young. 1994. Genetic dissection of oligogenic resistance to bacterial wilt in tomato. Mol Plant Microbe Interact 7: 464–471.

Daunay, M.C. H. Laterrot, J.W. Scott, P. Hanson and J-F Wang. 2010. Tomato resistance to bacterial wilt caused by Ralstonia solanacearum E.F. Smith: ancestry and peculiarities. Rpt Tomato Genetics Coop 60: 6–40.

Geethanjali, S., P. Kadirvel, R. de la Peña, E.S. Rao and J-F Wang. 2011. Development of tomato SSR markers from anchored BAC clones of chromosome 12 and their application for genetic diversity analysis and linkage mapping. Euphytica 178: 283–295.

Hanson, P.M., J.-F. Wang, O. Licardo, Hanindin, S.Y Mah, G.L. Hartman, Y.-C. Lin and J.-t. Chen. 1996. Variable reaction of tomato lines to bacterial wilt evaluated at several locations in Southeast Asia. HortScience 31: 143–146.

Hayward, A.C. 19991. Biology and epidemiology of bacterial wilt caused by Pseudomonas solanacearum. Annual Rev. Phytopathol. 143: 349–352

Scott, J.W., J.-F. Wang, and P.M. Hanson. 2005. Breeding tomatoes for resistance to bacterial wilt, a global view. Acta Hort 695: 161–172.

Wang, J-F, P. Hanson and J.A. Barnes. 1998. Worldwide evaluation of an international set of resistance sources to bacterial wilt in tomato. P. 269–275. In: P. Prior, C. Allen and J. Elphinstone (eds.) Bacterial Wilt Disease: Molecular and Ecological Aspects, Springer- Verlag, Berlin.Wang, J-F, F-I Ho, H.T.H Truong, S-M Huang, C.H. Balatero, V. Dittapongpitch and N. Hidayati. 2013. Identification of major QTLs associated with stable resistance of tomato cultivar ‘Hawaii 7996’ to Ralstonia solanacearum. Euphytica 190: 241–253.

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Distribution of major QTLs associated with resistance to Ralstonia solanacearum phylotype I strain in a global set of resistant tomato accessions

Fang-I Ho1, Chi-Yun Chung, and Jaw-Fen Wang1 1AVRDC – The World Vegetable Center, 60 Yi-Min Liao, Shanhua, Tainan, Taiwan

Introduction Bacterial wilt caused by Ralstonia solanacearum is one of the most devastating diseases of tomatoes, particularly in tropical and subtropical humid countries (Elphinstone, 2005). The bacterium displays a large genetic and phenotypic variation and has an exceptional ability to survive for a long time in water, soil, and the plant rhizosphere. The use of host resistance is the cheapest, most efficient, and most environmentally friendly method available for controlling the disease. However, breeding for stable resistance is challenging due to the location-specific and strain-specific nature of the resistance (Hanson et al., 1996; Lopes et al., 1994; Prior et al., 1990). Traditionally, R. solanacearum has been classified into five races and six biovars according to host ranges and biochemical properties, respectively (Denny, 2006). Recent genetic and phylogeny studies indicate the presence of four different phylotypes related to the geographical origin of the strains (Fegan and Prior, 2005). Genetic and phenotypic variation exist among strains of the same phylotype. For example, variation in virulence of phylotype I strains has been reported in Taiwan (Jaunet and Wang, 1999). The virulence was measured based on interactions of the pathogen and tomato varieties with different levels of resistance. Numerous tomato genotypes with resistance to bacterial wilt have been reported (Scott et al., 2005). Examination of the pedigrees and resistance sources used in major breeding programs worldwide showed the frequent exchange of plant genotypes among breeders (Daunay et al., 2010). Among the resistance sources used, Hawaii 7996 (Solanum lycopersicum) is one showing stable resistance (Wang et al., 1998). At least two major quantitative trait loci (QTL) associated with resistance in Hawaii 7996 have been identified (Thoquet et al., 1996a, b; Wang et al., 2000; Carmeille et al., 2006). Recently, the importance of the two major QTLs Bwr-12 and Bwr-6 contributing to the stable resistance of Hawaii 7996 was confirmed using a linkage map with good coverage and phenotype datasets collected against phylotype I and II strains under different environments (Wang et al., 2013). Bwr-12 was located in a 2.8-cM interval of chromosome 12; it controlled 17.9% to 56.1% of resistance variation against all phylotype I strains in five countries, but not against phylotype II strains. Bwr-6 on chromosome 6 explained 11.5% to 22.2% of the phenotypic variation against a few phylotype I strains and one phylotype II strain. The location of Bwr-6 differed with phenotype datasets and varied along a 15.5-cM region. Such results may be due to the effect of the environment on symptom expression and the interactions with different pathogen strains.

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Although Bwr-12 and Bwr-6 were associated with stable resistance to bacterial wilt in Hawaii 7996, it is not clear whether the same QTLs are present among the other sources of resistance, especially those showing stable resistance similar to Hawaii 7996. Therefore, the objectives of this study were to examine the distribution of Bwr-12 and Bwr-6 in a global set of resistant tomato accessions and to correlate their genotypes with disease reactions against phylotype I strains differing in virulence levels.

Materials and Methods Plant materials Tomato accessions resistant to bacterial wilt were selected based on the ancestry indicated by Daunay et al. (2010) and Lebeau et al. (2011). A total of 16 tomato accessions from different origins (Fig. 1) were used in this study. Tomato accession WVa700 was used as the susceptible control. Seeds were provided by AVRDC – The World Vegetable Center’s Genetic Resources and Seed Unit and Tomato Breeding group. Genotypes of SSR markers associated with Bwr-12 and Bwr-6 The simple sequence repeat (SSR) markers associated with Bwr-12 and Bwr-6 reported in Wang et al. (2013) were used in this study (Table 1). Genomic DNA of plants was purified from fresh leaves using GenEluteTM plant genomic DNA miniprep kit (Sigma, USA) following the instruction manual. PCR amplification of SSR markers was conducted in a reaction mixture consisting of 20 ng DNA, 0.3 μM of primer, 200 μM of deoxyribonucleotides, 50 mM KCl, 10 mM Tris HCl (pH 8.3), 1.5 mM MgCl2, and 0.5 unit of hot start Taq DNA polymerase. The temperature profile used for PCR amplification included initial denaturation at 94 °C for 10 min, 30 cycles of 94 °C for 30 s, 55 °C for 45 s, 72 °C for 45 s, followed by a final extension at 72 °C for 7 min. PCR products were analyzed on 6% non-denaturing polyacrylamide gel in TBE buffer. After electrophoresis, the gels were stained with ethidium bromide. Capillary electrophoresis was conducted to verify genotypes of SLM6-94, SLM6-118, and SLM6-136 due to the small size difference of amplicons derived from VC 8-1-2-1, VC 11-3-1-8, UCPA 1169, CLN 1463, Saturn, and CRA 84-26-3. Forward primers of these markers were labeled with fluorescent dye TMARA, HEX and FAM individually. Amplicons of the three markers were mixed with equal concentration. Capillary electrophoresis was performed on an ABI3730XL DNA analyzer (Applied Biosystems) by Genomics Biosci & Tech, Taiwan. Three kinds of alleles were noted for each marker, i.e. “H” (resistance allele), “W” (susceptibility allele), or “-“ (other allele), when the SSR marker showed the same size as Hawaii 7996, WVa700, or neither.

Bacterial strains and inoculation R. solanacearum strains isolated from tomato (Pss190, Pss4, and Pss216) were used. They were selected to represent strains with high, medium and low virulence, respectively

Page 23 of 64 RESEARCH REPORTS TGC REPORT VOLUME 63, 2013 according to Jaunet et al. (1999). They all belong to phylotype I, race 1 and biovar 3, except Pss190, which is a biovar 4 strain. Cultures of R. solanacearum were routinely grown (2 days, 30 °C) on 2,3,5-triphenyl tetrazolium chloride medium (Kelman, 1954). For inoculum preparation, the bacteria were multiplied on 523 medium (Kado and Heskett, 1970) at 30 °C for 24 h, and a bacterial suspension was prepared in sterilized distilled water and adjusted to 8 approximately 10 CFU per ml (OD600nm=0.3). The experiment was conducted following a randomized complete block design with three replications and ten plants per accession. Trials were conducted twice in a greenhouse. The ranges of mean temperature and relative humidity were 25 °C to 33 °C and 53% to 95% for Trial 1; for Trial 2, they were 20 °C to 29 °C and 69% to 99%. Three-week-old plants were inoculated by drenching the bacterial suspension (108 CFU/ml) on the soil surface near the base of plants at the five fully expanded leaf stage (Wang et al., 2000). Percentage of wilted plants per replication was recorded four weeks after inoculation. The data on percentage wilted plants was transformed to arcsine of its square root and analyzed using the PROC MIXED procedure of SAS (SAS v 9.2, SAS Institute, Cary, NC, USA). Tomato accessions were grouped according to the genotypes of SSR markers associated with Bwr-12 and Bwr-6 (Table 3). Group mean comparisons were performed using Tukey’s test (HSD) at P<0.05.

Results and discussion Bwr-12 and/or Bwr-6 were commonly presented in the 16 tested tomato accessions resistant to bacterial wilt (Table 2). Based on the genotypes of Bwr-12 and Bwr-6, the accessions could be grouped into three categories. Group A accessions were homozygous for the resistance allele of the two QTLs, Group B was homozygous for the Bwr-12 resistance allele and variable alleles of the Bwr-6 markers, while Group C was homozygous for the Bwr- 12 susceptibility allele and the resistance alleles of all 7 markers of Bwr-6. The presence of both Bwr-12 and Bwr-6 in accessions that originated from the University of North Carolina suggested these QTLs might have been utilized since the 1930s (Dauney et al., 2010). Bacterial wilt resistant lines bred in North Carolina were widely used by other breeding programs, and as a result, the two QTLs were commonly present in resistant accessions worldwide. Because the exact pedigrees of many resistant accessions, including Hawaii 7996, were not available, it is impossible to verify the exact sources of the two QTLs. The AVRDC breeding program has used UPCA116, Saturn, and CRA series as sources of resistance. These sources all possess Bwr- 12, which is present in most of the resistant lines bred at AVRDC (unpublished data). The disease reactions of the tested accessions were examined against three R. solanacearum strains. The disease pressure was higher in Trial 1, as evidenced by the higher wilting incidence resulting from the presence of higher temperature. The difference in virulence of the three R. solanacearum strains was more obvious in Trial 2 when the

Page 24 of 64 RESEARCH REPORTS TGC REPORT VOLUME 63, 2013 temperature was lower. The mean percentages of wilted plants over tested accessions caused by Pss190, Pss4, and Pss216, were 93.7%, 66.9%, and 68.6% in Trial 1, and 90.8%, 30.0%, and 17.5% in Trial 2. In order to examine the contribution of Bwr-6 and Bwr-12 to the resistance against different pathogen strains, a group mean comparison was conducted (Table 3). The ranking of the three groups was not consistent when interacting with different strains. Overall, Group A accessions, with the presence of both Bwr-6 and Bwr-12, showed a significantly lower percentage of wilted plants than the other groups, except when interacting with Pss190 and Pss216 under lower temperatures. When interacting with Pss4, Group B, possessing Bwr-12 only, had significantly lower disease incidence than Group C possessing Bwr-6 only. These results are similar to those reported by Wang et al. (2013), when examining the effects of Bwr-6 and Bwr-12 using recombinant inbred lines derived from Hawaii 7996. Significant differences in disease reactions between accessions within the same group were observed in Group A when interacting with Pss190 in both trials, in Group B when interacting with Pss4 and Pss216 in Trial 1, and in Group C when interacting with Pss190 and Pss216 in Trial 1 and with Pss4 in Trial 2. This implies the performance of QTLs could vary depending on the genetic backgrounds. The presence of other minor QTLs may cause this kind of variation. All the four accessions in Group A, namely Hawaii 7996, TML114, TML46, and R3034, were among the tomato entries showing the best stable resistance based on evaluations conducted in 11 countries (Wang, et al., 1998). Wang et al. (2013) indicated Bwr-12 contributes to resistance against phylotype I only, while Bwr-6 contributes to resistance against both phylotype I and II. Therefore, pyramiding the two major QTLs is essential when breeding for stable resistance to bacterial wilt. In conclusion, tomato accessions resistant to bacterial wilt from different origins possess Bwr-6 and/or Bwr-12. Presence of both Bwr-6 and Bwr-12 contributes to stable resistance against the phylotype I strain with different virulence levels. The fact that few tested resistant accessions have both QTLs indicates that pyramiding both QTLs could not be achieved efficiently using conventional disease screening. Marker-assisted selection should increase the efficiency. Bwr-12 markers have been used and found effective at AVRDC (Hanson et al., 2013 in this TGC Report). Fine-mapping of Bwr-6 is underway at AVRDC to develop useful markers for pyramiding the two QTLs.

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Table 1. Sequences of simple sequence repeat markers used in this study Product Marker name repeat motif Primer sequence (5'-3') size (bp) SLM 12-2 (TA)11 f: ATCTCATTCAACGCACACCA 209 r: AACGGTGGAAACTATTGAAAGG SLM 12-10 (AT) 21 f: ACCGCCCTAGCCATAAAGAC 242 r: TGCGTCGAAAATAGTTGCAT SLM 6-124 (TAT) 10 f: CATGGGTTAGCAGATGATTCAA 292 r: GCTAGGTTATTGGGCCAGAA SLM 6-118 (AAT) 18 f: TCCCAAAGTGCAATAGGACA 183 r: CACATAACATGGAGTTCGACAGA SLM 6-119 (AT) 24 f: GCCTGCCCTACAACAACATT 255 r: CGACATCAAACCTATGACTGGA SLM 6-136 (AT) 37 f: CCAGGCCACATAGAACTCAAG 290 r: ACAGGTCTCCATACGGCATC SLM 6-17 (TA) 12 f: TCCTTCAAATCTCCCATCAA 186 r: ACGAGCAATTGCAAGGAAAA SLM 6-94 (TA) 33 f: CTAAATTTAAATGGACAAGTAATAGCC 276 r: CACGATAGGTTGGTATTTTCTGG SLM 6-110 (ATT) 22 f: AGAATGCGGAGGTCTGAGAA 274 r: ATCCCACTGTCTTTCCACCA

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Table 2. Genotypes of simple sequence repeat markers associated with Bwr-12 and Bwr-6 in tested tomato accessions.

Bwr-12 Bwr-6a Bwr-6b Bwr-6c Bwr-6d

SLM SLM SLM SLM SLM SLM SLM SLM SLM Accession 12-2 12-10 6-124 6-118 6-119 6-136 6-17 6-94 6-110 Hawaii 7996 Ha H H H H H H H H TML114 H H H H H H H H H TML46 H H H H H H H H H R3034 H H H H H H H H H VC 8-1-2-1 H H W - - W - W - UCPA 1169 H H W - H - - H H CLN 1463A H H W - - W - - - Saturn H H W W - - - W - CRA 84-26-3 H H W - - W - - - VC 11-3-1-8 H H W ------BF-Okitsu W W H H H H H H H L285 W W H H H H H H H CRA 66 W W H H H H H H H NC72TR4-4 W W H H H H H H H IRAT L3 W W H H H H H H H Venus W W H H H H H H H WVa700 W W W W W W W W W (Sus.) a “H” means homozygous resistance allele as Hawaii 7996; “W” means homozygous susceptibility allele as WVa700; “-” means others alleles

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Figure 1. Origins and relationships among tomato accessions resistant to bacterial wilt (modified from Lebeau et al., 2011 and Daunay et al., 2010). The presence of Bwr-6 ( ), Bwr- 12 ( ) or both ( ) in the accession is highlighted according to the results of marker assays. Accessions without highlights were not tested.

Mulua (Guatemala) PI129080 () S. lycopersicum S. pimpinellifolium or S. lycopersicum var. cerasiforme

Univ. North Carolina PI127805A () S. pimpinellifolium NC 72 TR NC1953- Saturn Venu 4-4 64N s

Beltsville3814 Hawaii 7996 199 UPR S. lycopersicum var. pyriforme Univ. Hawaii

Univ. Puerto Rico

West indies landrace TML114 R3034 TML4 (tomadose) OTB2 4 6 UPCA116 VC8-1-2-1 VC11-3-1-8 BF-Okitsu 9 Univ. Philippines Hort. Res. Stn.

IRAT L3 CRA66 CRA84-26-3 CLN1463 L285 L3 S. lycopersicum IRAT INRA var. cerasiforme (Martinique) (Guadeloupe) AVRDC, Taiwan French West Indies Institutes

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Table 3. Disease incidence of tomato accessions having different genotypes of two major QTLs against phylotype I strains of Ralstonia solanacearum in two trials.

b b Bwr Bwr Trial 1 Trial 2 Accession -12 -6 Pss190 Pss4 Pss216 Pss190 Pss4 Pss216 c

Hawaii 7996 H H 80.0 26.7 36.7 86.7 3.3 10.0 a TML114 H H 83.3 70.2 A c 40.0 29.1 A 26.7 32.1 A 86.7 68.8 A 3.3 6.7 A 3.3 11.7 A TML46 H H 73.3 (82.5) 20.0 (25.0) 20.0 (29.2) 73.3 (85.0) 3.3 ( 3.3) 6.7 ( 7.5) Group A R3034 H H 93.3 13.3 33.3 93.3 3.3 10.0 VC 8-1-2-1 H W/- 100.0 66.7 91.7 100.0 3.3 23.3

UCPA 1169 H H/W/- 100.0 36.7 56.7 93.3 13.3 0.0 CLN 1463 H W/- 100.0 89.1 B 60.0 55.8 B 93.3 72.5 C 100.0 85.7 B 6.7 17.2 B 23.3 22.1 B Saturn H W/- 100.0 (100.0) 96.7 (63.9) 86.7 (86.4) 100.0 (97.8) 26.7 (13.9) 23.3 (17.8) Group B CRA 84-26-3 H W/- 100.0 56.7 96.7 93.3 10.0 16.7 VC 11-3-1-8 H W/- 100.0 66.7 93.3 100.0 23.3 20.0 BF-Okitsu W H 80.0 80.0 76.7 80.0 40.0 23.3 L285 W H 93.3 83.3 76.7 80.0 40.0 16.7 20.1 CRA 66 W H 96.7 81.2 B 100.0 78.9 C 46.7 59.2 B 83.3 70.1 A 46.7 45.7 C 3.3 AB

Group C NC72 TR4-4 W H 100.0 (93.9) 96.7 (92.2) 90.0 (7.1.7) 86.7 (87.2) 60.0 (55.0) 13.3 (17.2) IRAT L3 W H 93.3 96.7 80.0 96.7 73.3 33.3 Venus W H 100.0 96.7 60.0 96.7 70.0 13.3

WVa700 89.1 B 89.1 C 89.1 C 77.4 A 69.8 D 49.2 C W W 100.0 100.0 100.0 93.3 83.3 56.7 (Sus.) (100.0) (100.0) (100.0) (93.3) (83.3) (56.7) Group D

a Tomato accessions were grouped based on genotypes of Bwr-12 and Bwr-6. The genotypes were noted as “H”, homozygous for the Hawaii7996 allele, “W” homozygous for the WVa700 allele, or “ –“ for the other alleles. b The two trials were conducted in different seasons. The mean temperature and relative humidity range was 25°C to 33°C and 53% to 95% for Trial 1, and 20°C to 29°C and 63% to 99% for Trial 2. c Percentage of wilted plants was transformed to arcsine of the square root for analysis of variance. Transformed data are used for column-wise mean comparisons. The actual means are presented in parentheses. The group means for each strain were compared using Tukey’s test (HSD) at P<0.05. Comparison of means within and across groups was based on Least Significant Difference (LSD) at the P<0.05. The LSD0.05 values for Pss190, Pss4, and Pss216 in Trial 1 are 13.3%, 23.3% and 20.0%, respectively. For Trial 2, the LSD values are 16.7%, 20.0%, and 30.0% for Pss190, Pss4 and Pss216, respectively.

Page 29 of 64 RESEARCH REPORTS TGC REPORT VOLUME 63, 2013

Reference Denny T.P. 2006. Plant Pathogenic Ralstonia species. In: Plant-Associated Bacteria. Gnanamanickam SS (ed.) Springer, Dordrecht, the Netherlands, pp 573-644. Daunay M.C., Laterrot H., Scott J.W., Hanson P., Wang J.F. 2010. Tomato resistance to bacterial wilt caused by Ralstonia solanacearum E.F. Smith: ancestry and peculiarities. Rept. Tomato Genet. Coop. 60:6-40. Elphinstone J.G. 2005. The current bacterial wilt situation: A global overview. In: Bacterial Wilt Disease and the Ralstonia solanacearum species complex. Allen C, Prior P, and Hayward A.C. (eds) APS Press. St. Paul, USA pp 9-28. Fegan M., Prior P., 2005. How complex is the "Ralstonia solanacearum species complex". In: Bacterial wilt disease and the Ralstonia solanacearum species complex. Allen C., Prior P., and Hayward C. (eds) APS Press, St. Paul, USA pp 449-461. Hanson P., Wang J.F., Licardo O., Hanudin, Mah S.Y., Hartman G.L., Lin Y.C., Chen J.T. 1996. Variable reaction of tomato lines to bacterial wilt evaluated at several locations in Southeast Asia. Hortscience 31:143-146. Jaunet T. and Wang J.F. 1999. Variation in genotype and aggressiveness diversity of Ralstonia solanacearum race 1 isolated from tomato in Taiwan. Phytopathology 89:320-327. Lopes C.A., Quezado Soares A.M., De Melo P.E. 1994. Differential resistance of tomato cultigens to biovars I and III of Pseudomonas solanacearum. Plant Dis. 78: 1091-1094. Prior P., Steva H., Cadet, P. 1990. Aggressiveness of strains of Pseudomonas solanacearum from the French West Indies (Martinique and Guadeloupe) on tomato. Plant Dis 74:962- 965. Scott J.W., Wang J.F., and Hanson P. 2005. Breeding tomatoes for resistance to bacterial wilt, a Global view. Acta Hort 695:161-172. Wang J.F., Hanson P., and Barnes J.A. 1998. Worldwide evaluation of an international set of resistant sources to bacterial wilt in tomato. In: Bacterial wilt disease-Molecular and ecological aspects. Prior P., Allen C., Elphinstone J.G. (eds.) Springer-Verlag, Berlin pp269- 275. Wang J.-F., Ho F.-I., Truong H.T.H., Huang S.-M., Balatero C.H., Dittapongpitch V., Hidayati N. 2013. Identification of major QTLs associated with stable resistance of tomato cultivar ‘Hawaii 7996’ to Ralstonia solanacearum. Euphytica 190:241-252.

Page 30 of 64 VARIETAL PEDIGREES TGC REPORT VOLUME 63, 2013

Varietal Pedigrees

Scott, J.W. 2013. Fla. 8111B a large-fruited, globe shaped tomato breeding line Pedigree:

Characteristics: Fruit: Large, globe shaped, smooth, firm, light-green shoulders, moderate crack and check resistance, susceptible to stem scar water uptake, graywall resistant Plant: sp, I, I-2, Ve, Sm, large somewhat open vine, very susceptible to bacterial spot Utility and maturity: Fresh market inbred, mid-season maturity, good combiner in F1 hybrids

VARIETAL PEDIGREES TGC REPORT VOLUME 63, 2013

Scott, J.W. and S.F. Hutton. 2013. Fla. 8233, a germplasm line with partial resistance to bacterial spot races T1, T2, T3, T4 and Xanthomonas gardneri. Pedigree:

Characteristics: Fruit: Medium size, flat round some with irregular shape, stellate to irregular blossom scars sometimes with holes, firm, light-green shoulders, good crack resistance, some zippering Plant: sp, I, I-2, Sm, Rx-4, moderate vine size with dark green foliage, tendency for non-blighting when infected by foliar pathogens Utility and maturity: Donor inbred with moderate heat-tolerant fruit setting for development of improved fresh market inbreds with bacterial spot resistance. Partial resistance to bacterial spot races T1, T2, T3 (hypersensitive resistance), T4, and Xanthomonas gardneri, Rx-4 linked in cis with a QTL on chromosome 11 that provides broad spectrum bacterial spot resistance.

STOCK LIST TGC REPORT VOLUME 63, 2013

Revised List of Wild Species Stocks

Chetelat, R. T.

C.M. Rick Tomato Genetics Resource Center, Dept. of Plant Sciences, University of California, One Shields Ave., Davis, CA 95616

The following list of 1,153 accessions of wild tomatoes and allied Solanum species is a revision of the list published in TGC vol. 60, 2010. Other types of TGRC stocks are catalogued in TGC 61 (monogenic mutants) and TGC 62 (miscellaneous stocks). Accessions no longer available for distribution have been dropped from this list. New items include 34 previously inactive, newly regenerated accessions (designated herein with an asterisk after the LA number). Most of these wild species collections were never grown at Davis, either because other populations from similar locations were already being maintained and resources were limited, or because they were simply overlooked until now. Several of these ‘rescued’ accessions are noteworthy and interesting. Two S. habrochaites populations from (LA2859, LA2868) were regenerated, one of which is the first collection of this species from the southern coastal region (Dept. El Oro). An accession of S. habrochaites (LA1986) from the underrepresented Rio Moche drainage of Peru is noteworthy for its unusually vigorous growth and large flowers and fruit. New S. pennellii accessions include a collection from the Rio Casma (LA1773), only the second from that valley, and which displays the pedicel articulation in the ‘mid’ rather than the more common basal position. A new population of S. chilense (LA1931) from the underrepresented Rio Acari drainage showed the distinctive morphology of other accessions from the region. We also grew a S. chilense from Quebrada Socaire, notable for being near the southeastern geograhic limit of this species’ distribution. Several S. pimpinellifolium collections from the Lambayeque and regions in northern Peru were grown. We were especially happy to rescue these collections because many of the S. pimpinellifolium populations have been eliminated in this region of Peru due to intensification of agricultural practices, including use of sugarcane monocultures and widespread use of herbicides. Seed samples will be provided, upon request, for research, breeding or educational purposes. In most cases we provide 25 seed per accession for the self‐pollinated accessions, 50 for the outcrossing accessions, and 5‐10 for the allied Solanum species (S. juglandifolium, S. lycopersicoides, S. ochranthum and S. sitiens). These seed samples are meant to enable researchers to multiply seed for their future needs. NB: some accessions on this list may be temporarily unavailable for distribution during seed multiplication. The following list is sorted by species name, using the classification system of Peralta et al. (2008)1. More detailed information on each accession, including the collectors, field notes, geographic coordinates, images, etc, is available on our website, http://tgrc.ucdavis.edu.

1 Peralta, I. E., D. M. Spooner, S. Knapp (2008) Taxonomy of wild tomatoes and their relatives (Solanum sect. Lycopersicoides, sect. Juglandifolia, and sect. Lycopersicon; Solanaceae). Systematic Botany Monographs 84: 1-186.

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Solanum arcanum (L. peruvianum or L. peruvianum var. humifusum LA0378 Cascas Cajamarca Peru LA0385 San Juan (Rio Jequetepeque) Cajamarca Peru LA0389 Abra Gavilan Cajamarca Peru LA0392 Llallan Cajamarca Peru LA0441 Cerro Campana La Libertad Peru LA1027 Chilete Cajamarca Peru LA1031 Balsas Amazonas Peru LA1032 Aricapampa La Libertad Peru LA1346 Casmiche La Libertad Peru LA1350 Chauna Cajamarca Peru LA1351 Rupe Cajamarca Peru LA1394 Balsas ‐ Rio Utcubamba Amazonas Peru LA1395 Chachapoyas Amazonas Peru LA1396 Balsas (Chachapoyas) Amazonas Peru LA1626 Mouth of Rio Rupac Ancash Peru LA1708 Chamaya to Jaen Cajamarca Peru LA1984 Otuzco La Libertad Peru LA1985 Casmiche La Libertad Peru LA2150 Puente Muyuna Cajamarca Peru LA2151 Morochupa Cajamarca Peru LA2152 San Juan #1 Cajamarca Peru LA2153 San Juan #2 Cajamarca Peru LA2157 Tunel Chotano Cajamarca Peru LA2163 to Yamaluc Cajamarca Peru LA2164 Yamaluc Cajamarca Peru LA2172 Cuyca Cajamarca Peru LA2185 Pongo de Rentema Amazonas Peru LA2326 Above Balsas Amazonas Peru LA2327 Aguas Calientes Cajamarca Peru LA2328 Aricapampa La Libertad Peru LA2330 Chagual La Libertad Peru LA2331 Agallpampa La Libertad Peru LA2333 Casmiche La Libertad Peru LA2334 San Juan Cajamarca Peru LA2388 Cochabamba to Huambos (Chota) Cajamarca Peru LA2548 La Moyuna Cajamarca Peru LA2550 El Tingo, Chorpampa‐San Juan Cajamarca Peru LA2553 Balconcillo de San Marcos Cajamarca Peru LA2555 Marical ‐ Castilla La Libertad Peru LA2565 Potrero de Panacocha a Llamellin Ancash Peru LA2566* Cullachaca Ancash Peru LA2582 San Juan Cajamarca Peru LA2583 (autotetraploid) Peru LA2813* San Juan Cajamarca Peru LA2917 Chullchaca Ancash Peru

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Solanum cheesmaniae (L. cheesmanii) LA0166 Santa Cruz: Barranco, N of Punta Galapagos Ecuador LA0421 San Cristobal: cliff East of Wreck Bay Galapagos Ecuador LA0422 San Cristobal: Wreck Bay, Puerto Galapagos Ecuador LA0428 Santa Cruz: Trail Bellavista to Miconia Galapagos Ecuador LA0429 Santa Cruz: Crater in highlands Galapagos Ecuador LA0434 Santa Cruz: Rambech Trail Galapagos Ecuador LA0437 Isabela: Ponds North of Villamil Galapagos Ecuador LA0521 Fernandina: Inside Crater Galapagos Ecuador LA0522 Fernandina: Outer slopes Galapagos Ecuador LA0524 Isabela: Punta Essex Galapagos Ecuador LA0528B Santa Cruz: Academy Bay Galapagos Ecuador LA0529 Fernandina: Crater Galapagos Ecuador LA0531 Baltra: Barranco slope, N side Galapagos Ecuador LA0746 Isabela: Punta Essex Galapagos Ecuador LA0749 Fernandina: North side Galapagos Ecuador LA0927 Santa Cruz: Academy Bay Galapagos Ecuador LA0932 Isabela: Tagus Cove Galapagos Ecuador LA1035 Fernandina: Low elevation Galapagos Ecuador LA1036 Isabela: far north end Galapagos Ecuador LA1037 Isabela: Alcedo crater Galapagos Ecuador LA1039 Isabela: Cape Berkeley Galapagos Ecuador LA1040 San Cristobal: Caleta Tortuga Galapagos Ecuador LA1041 Santa Cruz: El Cascajo Galapagos Ecuador LA1042 Isabela: Cerro Santo Tomas Galapagos Ecuador LA1043 Isabela: Cerro Santo Tomas Galapagos Ecuador LA1138 Isabela: E of Cerro Azul Galapagos Ecuador LA1139 Isabela: W of Cerro Azul Galapagos Ecuador LA1402 Fernandina: W of Punta Espinoza Galapagos Ecuador LA1404 Fernandina: W flank caldera Galapagos Ecuador LA1406 Fernandina: SW rim caldera Galapagos Ecuador LA1407 Fernandina: caldera, NW bench Galapagos Ecuador LA1409 Isabela: Punta Albemarle Galapagos Ecuador LA1412 San Cristobal: opposite Isla Lobos Galapagos Ecuador LA1414 Isabela: Cerro Azul Galapagos Ecuador LA1427 Fernandina: WSW rim of caldera Galapagos Ecuador LA1447 Santa Cruz: Charles Darwin Station‐ Galapagos Ecuador LA1448 Santa Cruz: Puerto Ayora, Pelican Bay Galapagos Ecuador LA1449 Santa Cruz: Charles Darwin Station, Galapagos Ecuador LA1450 Isabela: Bahia San Pedro Galapagos Ecuador LA3124 Santa Fe: near E landing Galapagos Ecuador

Solanum chilense (L. chilense) LA0130 Moquegua Peru LA0294 Tacna Peru LA0456 Clemesi Moquegua Peru

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Solanum chilense (L. chilense) LA0458 Tacna Tacna Peru LA0460 Palca Tacna Peru LA0470 Taltal Antofagasta LA1029 Moquegua Moquegua Peru LA1030 N of Tacna Tacna Peru LA1782 Quebrada de Acari Peru LA1917 Llauta Peru LA1930 Quebrada Calapampa Arequipa Peru LA1931* Mina Santa Rosa Arequipa Peru LA1932 Minas de Acari Arequipa Peru LA1938 Quebrada Salsipuedes Arequipa Peru LA1958 Pampa de la Clemesi Moquegua Peru LA1959 Huaico Moquegua Moquegua Peru LA1960 Rio Osmore Moquegua Peru LA1961 Toquepala Tacna Peru LA1962* Huaico Tacna Tacna Peru LA1963 Rio Caplina Tacna Peru LA1965 Causuri Tacna Peru LA1967 Pachia, Rio Caplina Tacna Peru LA1968 Cause Seco, Tacna to Tarata Tacna Peru LA1969 Estique Pampa Tacna Peru LA1970 Tarata Tacna Peru LA1971 Palquilla Tacna Peru LA1972 Rio Sama Tacna Peru LA2404 Arica to Tignamar Arica‐Parinacota Chile LA2405 Tignamar Arica‐Parinacota Chile LA2406 Arica to Putre Arica‐Parinacota Chile LA2729* Pacahua quebrada Tarapaca Chile LA2731 Moquella Tarapaca Chile LA2737 Yala‐yala Tarapaca Chile LA2739 Crossroads Nama to Camina Tarapaca Chile LA2743* La Puntilla Arica‐Parinacota Chile LA2746 Asentamiento‐18 Arica‐Parinacota Chile LA2747 Alta Azapa Arica‐Parinacota Chile LA2748 Soledad Tarapaca Chile LA2749 Mina La Buena Esperanza Antofagasta Chile LA2750 Mina La Despreciada Antofagasta Chile LA2751 Pachica (Rio Tarapaca) Tarapaca Chile LA2753 Laonzana Tarapaca Chile LA2754 W of Chusmisa Tarapaca Chile LA2755 Banos de Chusmisa Tarapaca Chile LA2757 W of Chusmisa Tarapaca Chile LA2759 Mamina Tarapaca Chile LA2762 Quebradas de Mamina a Parca Tarapaca Chile LA2764 Codpa Arica‐Parinacota Chile

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Solanum chilense (L. chilense) LA2765 Timar Arica‐Parinacota Chile LA2767 Chitita Arica‐Parinacota Chile LA2768 Empalme Codpa Arica‐Parinacota Chile LA2771 Above Poconchile Arica‐Parinacota Chile LA2773 Zapahuira Arica‐Parinacota Chile LA2774 Socorama Arica‐Parinacota Chile LA2778 Chapiquina Arica‐Parinacota Chile LA2779 Cimentario Belen Arica‐Parinacota Chile LA2780 Belen to Lupica Arica‐Parinacota Chile LA2879 Peine Antofagasta Chile LA2880 Quebrada Tilopozo Antofagasta Chile LA2881* Quebrada Socaire Antofagasta Chile LA2882 Camar Antofagasta Chile LA2884 Ayaviri Antofagasta Chile LA2887 Quebrada Bandurria Antofagasta Chile LA2888 Loma Paposo Antofagasta Chile LA2891 Quebrada Taltal Antofagasta Chile LA2930 Quebrada Taltal Antofagasta Chile LA2931 Guatacondo Tarapaca Chile LA2932 Quebrada Gatico, Mina Escalera Antofagasta Chile LA2946 Guatacondo Tarapaca Chile LA2947* Guatacondo Tarapaca Chile LA2949 Chusmisa Tarapaca Chile LA2952 Camiña Tarapaca Chile LA2955 Quistagama‐Quisama Tarapaca Chile LA2957* Pozo Tarapaca Chile LA2965* Pachia Tacna Peru LA2980 Yacango Moquegua Peru LA2981A Torata Moquegua Peru LA3111 Tarata Tacna Peru LA3112 Estique Pampa Tacna Peru LA3113 Apacheta Tacna Peru LA3114 Quilla Tacna Peru LA3115 W of Quilla Tacna Peru LA3153 Desvio Omate (Rio de Osmore) Moquegua Peru LA3155 Quinistaquillas Moquegua Peru LA4106 Taltal Antofagasta Chile LA4107 Catarata Taltal Antofagasta Chile LA4108 Caleta Punta Grande Antofagasta Chile LA4109 Quebrada Canas Antofagasta Chile LA4117A San Pedro ‐ Paso Jama Antofagasta Chile LA4117B San Pedro ‐ Paso Jama Antofagasta Chile LA4118 Toconao Antofagasta Chile LA4119 Socaire Antofagasta Chile LA4120 Cahuisa Tarapaca Chile

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Solanum chilense (L. chilense) LA4121 Pachica ‐ Poroma Tarapaca Chile LA4122 Chiapa Tarapaca Chile LA4127 Alto Umayani Arica‐Parinacota Chile LA4129 Pachica (Rio Camarones) Arica‐Parinacota Chile LA4132 Esquina Arica‐Parinacota Chile LA4319 Alto Rio Lluta Arica‐Parinacota Chile LA4321 Quebrada Cardones Arica‐Parinacota Chile LA4324 Estacion Puquio Arica‐Parinacota Chile LA4327 Pachica, Rio Camarones Arica‐Parinacota Chile LA4329 Puente del Diablo, Rio Salado Antofagasta Chile LA4330 Caspana Antofagasta Chile LA4332 Rio Grande Antofagasta Chile LA4334 Quebrada Sicipo Antofagasta Chile LA4335 Quebrada Tucuraro Antofagasta Chile LA4336 Quebrada Cascabeles Antofagasta Chile LA4337 Quebrada Paposo Antofagasta Chile LA4338 Quebrada Taltal, Estacion Breas Antofagasta Chile LA4339 Quebrada Los Zanjones Antofagasta Chile

Solanum chmielewskii (L. chmielewskii) LA1028 Casinchihua Apurimac Peru LA1306 Ayacucho Peru LA1316 Ocros Ayacucho Peru LA1317 Pajonal Ayacucho Peru LA1318 Auquibamba Apurimac Peru LA1325 Puente Cunyac Apurimac Peru LA1327 Sorocata Apurimac Peru LA1330 Chalhuanca Apurimac Peru LA2639B Puente Cunyac Apurimac Peru LA2663 Tujtohaiya Cusco Peru LA2677 Huaypachaca #1 Cusco Peru LA2678 Huaypachaca #2 Cusco Peru LA2679 Huaypachaca #3 Cusco Peru LA2680 Puente Apurimac #1 Cusco Peru LA2681 Puente Apurimac #2 Cusco Peru LA2695 Chihuanpampa Cusco Peru

Solanum corneliomulleri (L. peruvianum or L. peruv. f. gladulosum) LA0103 , Rio Rimac Peru LA0107 Hacienda San Isidro, Rio Canete Lima Peru LA0364 Canta Lima Peru LA0366 12 Km W of Canta Lima Peru LA0444 Chincha #1 Ica Peru LA0451 Arequipa Arequipa Peru LA1133 Huachipa Lima Peru

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Solanum corneliomulleri (L. peruvianum or L. peruv. f. gladulosum) LA1271 Horcon Lima Peru LA1274 Pacaibamba Lima Peru LA1281 Sisacaya Lima Peru LA1283 Santa Cruz de Laya Lima Peru LA1284 Espiritu Santo Lima Peru LA1292 San Mateo Lima Peru LA1293 Matucana Lima Peru LA1294 Surco Lima Peru LA1296 Tornamesa Lima Peru LA1300 Santa Rosa de Quives Lima Peru LA1304 Pampano Peru LA1305 Ticrapo Huancavelica Peru LA1331 Nazca Ica Peru LA1339 Capillucas Lima Peru LA1369 San Geronimo Lima Peru LA1373 Asia Lima Peru LA1377 Navan Lima Peru LA1379 Caujul Lima Peru LA1473 Callahuanca, Santa Eulalia valley Lima Peru LA1551 Rimac Valley, Km 71 Lima Peru LA1552 Rimac Valley, Km 93 Lima Peru LA1554 Huaral to , Rio Chancay Lima Peru LA1609 Asia ‐ El Pinon Lima Peru LA1646 Yaso Lima Peru LA1647 Huadquina, Topara Ica Peru LA1653 Uchumayo, Arequipa Arequipa Peru LA1677 Fundo Huadquina, Topara Ica Peru LA1694 Cacachuhuasiin, Cacra Lima Peru LA1722 Ticrapo Viejo Huancavelica Peru LA1723 La Quinga Ica Peru LA1744 Putinza Lima Peru LA1910 Tambillo Huancavelica Peru LA1937 Quebrada Torrecillas Arequipa Peru LA1944 Rio Atico Arequipa Peru LA1945 Caraveli Arequipa Peru LA1973 Yura Arequipa Peru LA2717 Chilca Lima Peru LA2721 Putinza Lima Peru LA2724 Huaynilla Lima Peru LA2962 Echancay Arequipa Peru LA2981B Torata Moquegua Peru LA3154 Otora‐Puente Jaguay Moquegua Peru LA3156 Omate Valley Moquegua Peru LA3157* Quebrada Tinajas Lima Peru LA3219 Catarindo Arequipa Peru

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Solanum galapagense (L. cheesmanii f. minor) LA0317 Bartolome Galapagos Ecuador LA0426 Bartolome: E of landing Galapagos Ecuador LA0436 Isabela: Villamil Galapagos Ecuador LA0438 Isabela: coast at Villamil Galapagos Ecuador LA0480A Isabela: Cowley Bay Galapagos Ecuador LA0483 Fernandina: inside crater Galapagos Ecuador LA0526 Pinta: W Side Galapagos Ecuador LA0527 Bartolome: W side, Tower Bay Galapagos Ecuador LA0528 Santa Cruz: Academy Bay Galapagos Ecuador LA0530 Fernandina: crater Galapagos Ecuador LA0532 Pinzon: NW side Galapagos Ecuador LA0747 : Cape Trenton Galapagos Ecuador LA0748 Santiago: E Trenton Islet Galapagos Ecuador LA0929 Isabela: Punta Flores Galapagos Ecuador LA0930 Isabela: Punta Tortuga Galapagos Ecuador LA1044 Bartolome Galapagos Ecuador LA1136 Gardner‐near‐Floreana Islet Galapagos Ecuador LA1137 Rabida: N side Galapagos Ecuador LA1141 Santiago: N crater Galapagos Ecuador LA1400 Isabela: N of Punta Tortuga Galapagos Ecuador LA1401 Isabela: N of Punta Tortuga Galapagos Ecuador LA1403 Fernandina: W of Punta Espinoza Galapagos Ecuador LA1408 Isabela: SW volcano, Cape Berkeley Galapagos Ecuador LA1410 Isabela: Punta Ecuador Galapagos Ecuador LA1411 Santiago: N James Bay Galapagos Ecuador LA1452 Isabela: E slope, Volcan Alcedo Galapagos Ecuador LA1508 Floreana: Corona del Diablo Islet Galapagos Ecuador LA1627 Isabela: Darwin's Lake Galapagos Ecuador

Solanum habrochaites (L. hirsutum or L. hirsutum f. glabratum) LA0094 Canta‐Yangas Lima Peru LA0361 Canta Lima Peru LA0386 Cajamarca Cajamarca Peru LA0387 Santa Apolonia Cajamarca Peru LA0407 El Mirador, Guayaquil Guayas Ecuador LA1033 Hacienda Taulis Lambayeque Peru LA1223 Alausi Chimborazo Ecuador LA1252 Loja, Jardin Botanico Loja Ecuador LA1253 Pueblo Nuevo ‐ Landangue Loja Ecuador LA1255 Loja Loja Ecuador LA1264 Bucay Chimborazo Ecuador LA1265 Rio Chimbo Chimborazo Ecuador LA1266 Pallatanga Chimborazo Ecuador LA1295 Surco Lima Peru LA1298 Yaso Lima Peru

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Solanum habrochaites (L. hirsutum or L. hirsutum f. glabratum) LA1347 Empalme Otusco La Libertad Peru LA1352 Rupe Cajamarca Peru LA1353 Contumaza Cajamarca Peru LA1354 Contumaza to Cascas Cajamarca Peru LA1361 Pariacoto Ancash Peru LA1362 Chacchan Ancash Peru LA1363 Alta Fortaleza Ancash Peru LA1366 Cajacay Ancash Peru LA1378 Navan Lima Peru LA1391 Bagua to Olmos Cajamarca Peru LA1392 ‐ Casma Road Ancash Peru LA1393 Huaraz ‐ Caraz Ancash Peru LA1557 Huaral to Cerro de Pasco, Rio Chancay Lima Peru LA1559 Desvio Huamantanga‐Canta Lima Peru LA1560 Matucana Lima Peru LA1624 Jipijapa Manabi Ecuador LA1625 S of Jipijapa Manabi Ecuador LA1648 Above Yaso Lima Peru LA1681 Mushka Lima Peru LA1691 Yauyos Lima Peru LA1695 Cacachuhuasiin, Cacra Lima Peru LA1696 Huanchuy to Cacra Lima Peru LA1717 Sopalache Peru LA1718 Huancabamba Piura Peru LA1721 Ticrapo Viejo Huancavelica Peru LA1731 Rio San Juan Huancavelica Peru LA1736 Pucutay Piura Peru LA1737 Cashacoto Piura Peru LA1738 Desfiladero Piura Peru LA1739 Canchaque to Cerran Piura Peru LA1740 Huancabamba Piura Peru LA1741 Sondorillo Piura Peru LA1753 Surco Lima Peru LA1761 Rio Chillon Lima Peru LA1764 West of Canta Lima Peru LA1772 West of Canta Lima Peru LA1775 Rio Casma Ancash Peru LA1777 Rio Casma Ancash Peru LA1778 Rio Casma Ancash Peru LA1779 Rio Casma Ancash Peru LA1918 Llauta Ayacucho Peru LA1927 Ocobamba Ayacucho Peru LA1928 Ocana Ayacucho Peru LA1978 Colca Ancash Peru LA1986* Casmiche La Libertad Peru

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Solanum habrochaites (L. hirsutum or L. hirsutum f. glabratum) LA2092 Chinuko Chimborazo Ecuador LA2098 Sabianga Loja Ecuador LA2099 Sabiango to Zozoranga Loja Ecuador LA2100 Sozoranga Loja Ecuador LA2101 San Pedro de Cariamanga Loja Ecuador LA2103 Lansaca Loja Ecuador LA2104 Pena Negra Loja Ecuador LA2105 Jardin Botanico, Loja Loja Ecuador LA2106 Yambra ‐ La Providencia Loja Ecuador LA2107 Los Lirios Loja Ecuador LA2108 Portete de Anganuma Loja Ecuador LA2109 Yangana #1 Loja Ecuador LA2110 Yangana #2 Loja Ecuador LA2114 San Juan Loja Ecuador LA2115 Pucala Loja Ecuador LA2116 Las Juntas Loja Ecuador LA2119 Saraguro Loja Ecuador LA2124 Cumbaratza Zamora‐Chinchipe Ecuador LA2128 Zumbi Zamora‐Chinchipe Ecuador LA2144 Chanchan Chimborazo Ecuador LA2155 Maydasbamba Cajamarca Peru LA2156 Ingenio Montan Cajamarca Peru LA2158 Rio Chotano Cajamarca Peru LA2159 Atonpampa Cajamarca Peru LA2167 Cementerio Cajamarca Cajamarca Peru LA2171 El Molino Piura Peru LA2174 Rio Chinchipe, San Augustin Cajamarca Peru LA2175 Timbaruca Cajamarca Peru LA2196 Caclic Amazonas Peru LA2204 Balsapata Amazonas Peru LA2314 San Francisco Amazonas Peru LA2321 Chirico Amazonas Peru LA2324 Leimebamba Amazonas Peru LA2329 Aricapampa La Libertad Peru LA2409 Rio Miraflores Lima Peru LA2541* Olmos‐Jaen Lambayeque Peru LA2552 Las Flores Cajamarca Peru LA2556 Puente Moche La Libertad Peru LA2567 Quita Ancash Peru LA2574 Cullaspungro Ancash Peru LA2648 Piura Peru LA2650 Ayabaca Piura Peru LA2651 Puente Tordopa Piura Peru LA2722 Puente Auco Lima Peru LA2812 Lambayeque Lambayeque Peru

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Solanum habrochaites (L. hirsutum or L. hirsutum f. glabratum) LA2855 Mollinomuna Loja Ecuador LA2859* Yangana Loja Ecuador LA2860 Cariamanga Loja Ecuador LA2861 Las Juntas Loja Ecuador LA2863 Macara Loja Ecuador LA2864 Sozorango Loja Ecuador LA2868* Arenillas El Oro Ecuador LA2869 Matala Loja Ecuador LA2975 Coltao Ancash Peru LA2976 Huangra Ancash Peru

Solanum huaylasense (L. peruvianum) LA0110 Cajacay Ancash Peru LA1358 Yautan Ancash Peru LA1360 Pariacoto Ancash Peru LA1364 Alta Fortaleza Ancash Peru LA1365 Caranquilloc Ancash Peru LA1979* Colca (Rio Fortaleza) Ancash Peru LA1981 Vocatoma Ancash Peru LA1982 Huallanca Ancash Peru LA1983 Rio Manta Ancash Peru LA2068 Chasquitambo Ancash Peru LA2561 Huallanca Ancash Peru LA2562 Canon del Pato Ancash Peru LA2563 Canon del Pato Ancash Peru LA2575 Valle de Casma Ancash Peru LA2808 Huaylas Ancash Peru LA2809 Huaylas Ancash Peru

Solanum juglandifolium LA2120 Sabanilla Zamora‐Chinchipe Ecuador LA2134 Tinajillas Zamora‐Chinchipe Ecuador LA2788 Quebrada La Buena Antioquia Colombia LA3322 San Juan ‐ Chiriboga Pichincha Ecuador LA3324 Sabanillas Zamora‐Chinchipe Ecuador LA3325 Cosanga Napo Ecuador

Solanum lycopersicoides LA1964 Chupapalca Tacna Peru LA1966 Palca Tacna Peru LA1990 Palca Tacna Peru LA1991 Causiri Tacna Peru LA2385 Chupapalca to Ingenio Tacna Peru LA2386 Chupapalca Tacna Peru LA2387 Lago Aricota Tacna Peru

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Solanum lycopersicoides LA2407 Arica to Putre Arica‐Parinacota Chile LA2408 Above Putre Arica‐Parinacota Chile LA2730 Moquella Tarapaca Chile LA2772 Zapahuira Arica‐Parinacota Chile LA2776 Catarata de Perquejeque Arica‐Parinacota Chile LA2777 Putre Arica‐Parinacota Chile LA2781 Desvio a Putre Arica‐Parinacota Chile LA2951 Quistagama Tarapaca Chile LA4123 Camina Tarapaca Chile LA4126 Camina ‐ Nama Tarapaca Chile LA4130 Pachica (Rio Camarones) Arica‐Parinacota Chile LA4131 Esquina Arica‐Parinacota Chile LA4320 Rio Lluta Arica‐Parinacota Chile LA4322 Quebrada Cardones Arica‐Parinacota Chile LA4323 Putre Arica‐Parinacota Chile LA4326 Cochiza, Rio Camarones Arica‐Parinacota Chile

Solanum lycopersicum (L. esculentum var. cerasiforme) LA0168 New Caledonia LA0292 Santa Cruz Galapagos Ecuador LA0349 (unknown origin) Unknown LA0384 Chilete (Rio Jequetepeque) Cajamarca Peru LA0475 Sucua Morona‐Santiago Ecuador LA0476 Sucua Morona‐Santiago Ecuador LA1025 Oahu: Wahiawa Hawaii USA LA1203 Ciudad Vieja Guatemala LA1204 Quetzaltenango Guatemala LA1205 Copan LA1206 Copan Honduras LA1207 LA1208 Sierra Nevada Colombia LA1209 Colombia LA1226 Sucua Morona‐Santiago Ecuador LA1227 Sucua Morona‐Santiago Ecuador LA1228 Macas, San Jacinto de los Monos Morona‐Santiago Ecuador LA1229 Macas Morona‐Santiago Ecuador LA1230 Macas Morona‐Santiago Ecuador LA1231 Tena Napo Ecuador LA1247 La Toma Loja Ecuador LA1268 Chaclacayo Lima Peru LA1286 San Martin de Pangoa Junin Peru LA1287 Fundo Ileana #1 Junin Peru LA1289 Fundo Ileana #3 Junin Peru LA1290 Mazamari Junin Peru LA1291 Satipo Granja Junin Peru

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Solanum lycopersicum (L. esculentum var. cerasiforme) LA1307 Hotel Oasis Ayacucho Peru LA1308 San Francisco Ayacucho Peru LA1310 Hacienda Santa Rosa Ayacucho Peru LA1311‐1 Santa Rosa Ayacucho Peru LA1311‐ Santa Rosa Puebla Ayacucho Peru LA1311‐ Santa Rosa Puebla Ayacucho Peru LA1311‐ Santa Rosa Puebla Ayacucho Peru LA1311‐ Santa Rosa Puebla Ayacucho Peru LA1311‐ Santa Rosa Puebla Ayacucho Peru LA1311‐ Santa Rosa Puebla Ayacucho Peru LA1311‐ Santa Rosa Puebla Ayacucho Peru LA1311‐ Santa Rosa Puebla Ayacucho Peru LA1311‐ Santa Rosa Puebla Ayacucho Peru LA1311‐ Santa Rosa Puebla Ayacucho Peru LA1311‐2 Santa Rosa Puebla Ayacucho Peru LA1311‐3 Santa Rosa Puebla Ayacucho Peru LA1311‐4 Santa Rosa Puebla Ayacucho Peru LA1311‐5 Santa Rosa Puebla Ayacucho Peru LA1311‐6 Santa Rosa Puebla Ayacucho Peru LA1311‐7 Santa Rosa Puebla Ayacucho Peru LA1311‐8 Santa Rosa Puebla Ayacucho Peru LA1311‐9 Santa Rosa Puebla Ayacucho Peru LA1312‐2 Paisanato Cusco Peru LA1312‐3 Paisanto Cusco Peru LA1312‐4 Paisanato Cusco Peru LA1314 Granja Pichari Cusco Peru LA1320 Hacienda Carmen Apurimac Peru LA1323 Pfacchayoc Cusco Peru LA1324 Hacienda Potrero, Quillabamba Cusco Peru LA1328 Rio Pachachaca Apurimac Peru LA1334 Pescadores Arequipa Peru LA1338 Puyo Napo Ecuador LA1372 Santa Eulalia Lima Peru LA1385 Quincemil Cusco Peru LA1386 Balsas Amazonas Peru LA1387 Quincemil Cusco Peru LA1388 San Ramon Junin Peru LA1423 Near Santo Domingo Pichincha Ecuador LA1429 La Estancilla Manabi Ecuador LA1453 Kauai: Poipu Hawaii USA LA1454 unknown Mexico LA1455 Gral Teran Nuevo Leon Mexico LA1456 Papantla Vera Cruz Mexico LA1457 Tehuacan Puebla Mexico LA1458 Huachinango Puebla Mexico

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Solanum lycopersicum (L. esculentum var. cerasiforme) LA1461 Los Banos Philippines LA1464 El Progreso, Yoro Honduras LA1465 Taladro, Comayagua Honduras LA1467 Cali Cauca Colombia LA1468 Fte. Casa, Cali Cauca Colombia LA1479 Sucua Morona‐Santiago Ecuador LA1480 Sucua Morona‐Santiago Ecuador LA1481 Sucua Morona‐Santiago Ecuador LA1482 Segamat Malaysia LA1483 Trujillo Saipan LA1509 Tawan Sabah, Borneo Malaysia LA1510 Mexico LA1511 Sete Lagoas Minas Gerais LA1512 Lago de Llopango El Salvador LA1519 Vitarte Lima Peru LA1540 Cali ‐ Popayan Cauca Colombia LA1542 Turrialba Costa Rica LA1543 Upper Parana Brazil LA1545 Becan Ruins Campeche Mexico LA1548 Fundo Liliana Junin Peru LA1549 Chontabamba Pasco Peru LA1569 Jalapa Vera Cruz Mexico LA1574 Nana Lima Peru LA1619 Pichanaki Junin Peru LA1620 Castro Alves Bahia Brazil LA1621 Rio Venados Hidalgo Mexico LA1622 Lusaka Zambia LA1623 Muna Yucatan Mexico LA1654 San Martin Peru LA1655 Tarapoto San Martin Peru LA1662 El Ejido Merida Venezuela LA1667 Cali Cauca Colombia LA1668 Acapulco Guerrero Mexico LA1673 Nana Lima Peru LA1701 Trujillo La Libertad Peru LA1705 Sinaloa Mexico LA1709 Desvio Yojoa Honduras LA1710 Cariare Limon Costa Rica LA1711 Zamorano Honduras LA1712 Pejibaye San Jose Costa Rica LA1713 CATIE, Turrialba Costa Rica LA1909 Quillabamba Cusco Peru LA1953 La Curva Arequipa Peru LA2076 Naranjitos LA2077 Paco, Coroica Bolivia

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Solanum lycopersicum (L. esculentum var. cerasiforme) LA2078 Mosardas Rio Grande de Sol Brazil LA2079 Maui: Kihei Hawaii USA LA2080 Maui: Kihei Hawaii USA LA2081 Maui: Kihei Hawaii USA LA2082 Arenal Valley Honduras LA2085 Kempton Park S. Africa LA2095 La Cidra, Olmedo Loja Ecuador LA2121 Yacuambi‐Guadalupe Zamora‐Chinchipe Ecuador LA2122A Yacuambi‐Guadalupe Zamora‐Chinchipe Ecuador LA2122B Yacuambi‐Guadalupe Zamora‐Chinchipe Ecuador LA2122C Yacuambi‐Guadalupe Zamora‐Chinchipe Ecuador LA2122D Yacuambi‐Guadalupe Zamora‐Chinchipe Ecuador LA2123A La Saquea Zamora‐Chinchipe Ecuador LA2123B La Saquea Zamora‐Chinchipe Ecuador LA2126A El Dorado Zamora‐Chinchipe Ecuador LA2126B El Dorado Zamora‐Chinchipe Ecuador LA2126C El Dorado Zamora‐Chinchipe Ecuador LA2126D El Dorado Zamora‐Chinchipe Ecuador LA2127 Zumbi Zamora‐Chinchipe Ecuador LA2129 San Roque Zamora‐Chinchipe Ecuador LA2130 Gualaquiza Zamora‐Chinchipe Ecuador LA2131 Bomboiza Zamora‐Chinchipe Ecuador LA2135 Limon Morona‐Santiago Ecuador LA2136 Bella Union Morona‐Santiago Ecuador LA2137 Tayusa Morona‐Santiago Ecuador LA2138A Chinimpimi Morona‐Santiago Ecuador LA2138B Chinimpimi Morona‐Santiago Ecuador LA2139A Logrono Morona‐Santiago Ecuador LA2139B Logrono Morona‐Santiago Ecuador LA2140A Huambi Morona‐Santiago Ecuador LA2140B Huambi Morona‐Santiago Ecuador LA2140C Huambi Morona‐Santiago Ecuador LA2141 Rio Blanco Morona‐Santiago Ecuador LA2142 Cambanaca Morona‐Santiago Ecuador LA2143 Nuevo Rosario Morona‐Santiago Ecuador LA2177A San Ignacio Cajamarca Peru LA2177B San Ignacio Cajamarca Peru LA2177C San Ignacio Cajamarca Peru LA2177E San Ignacio Cajamarca Peru LA2177F San Ignacio Cajamarca Peru LA2205A Santa Rosa de Mirador San Martin Peru LA2205B Santa Rosa de Mirador San Martin Peru LA2308 San Francisco San Martin Peru LA2312 Jumbilla #1 Amazonas Peru LA2313 Jumbilla #2 Amazonas Peru

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Solanum lycopersicum (L. esculentum var. cerasiforme) LA2392 Jakarta? Indonesia LA2393 Mercedes Guanacaste Costa Rica LA2394 San Rafael de Hojancha Guanacaste Costa Rica LA2411 Yanamayo Peru LA2587 LA2616 Naranjillo Huanuco Peru LA2617 El Oropel Huanuco Peru LA2618 Santa Lucia, Tulumayo Huanuco Peru LA2619 Caseria San Augustin Loreto Peru LA2620 La Divisoria Loreto Peru LA2621 3 de Octubre Loreto Peru LA2624 Umashbamba Cusco Peru LA2625 Chilcachaca Cusco Peru LA2626 Pacchac‐chico Cusco Peru LA2627 Pacchac‐chico Cusco Peru LA2629 Echarate/Tornillochayoc Cusco Peru LA2630 Calzada Cusco Peru LA2631 Chontachayoc Cusco Peru LA2632 Maranura Cusco Peru LA2633 Huayopata Cusco Peru LA2635 Huayopata Cusco Peru LA2636 Sicre Cusco Peru LA2637 Sicre Cusco Peru LA2640 Molinopata Apurimac Peru LA2642 Molinopata Apurimac Peru LA2643 Bella Vista Apurimac Peru LA2660 San Ignacio de Moxos Beni Bolivia LA2664 Yanahuaya Puno Peru LA2667 Pajchani Puno Peru LA2668 Cruz Playa Puno Peru LA2669 Huayvaruni #1 Puno Peru LA2670 Huayvaruni #2 Puno Peru LA2671 San Juan del Oro, School Puno Peru LA2673 Chuntopata Puno Peru LA2674 Huairurune Puno Peru LA2675 Casahuiri Puno Peru LA2683 Consuelo Cusco Peru LA2684 Patria Cusco Peru LA2685 Gavitana Madre de Dios Peru LA2686 Yunguyo Madre de Dios Peru LA2687 Mansilla Madre de Dios Peru LA2688 Santa Cruz near Shintuyo #1 Madre de Dios Peru LA2689 Santa Cruz near Shintuyo #2 Madre de Dios Peru LA2690 Atalaya Cusco Peru LA2691 Rio Pilcopata Cusco Peru

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Solanum lycopersicum (L. esculentum var. cerasiforme) LA2692 Pilcopata #1 Cusco Peru LA2693 Pilcopata #2 Cusco Peru LA2694 Aguasantas Cusco Peru LA2700 Aoti, Satipo Junin Peru LA2702 Kandy #1 Sri Lanka LA2709 Bidadi, Bangalore Karnataka India LA2710 Porto Firme Brazil LA2782 El Volcan #1 ‐ Pajarito Antioquia Colombia LA2783 El Volcan #2 ‐ Titiribi Antioquia Colombia LA2784 La Queronte, Antioquia Colombia LA2785 El Bosque, Andes Antioquia Colombia LA2786 Andes #1 Antioquia Colombia LA2787 Andes #2 Antioquia Colombia LA2789 Canaveral Antioquia Colombia LA2790 Buenos Aires Antioquia Colombia LA2791 Rio Frio Antioquia Colombia LA2792 Tamesis Antioquia Colombia LA2793 La Mesa Antioquia Colombia LA2794 El Libano Antioquia Colombia LA2795 Camilo Antioquia Colombia LA2807 Taypiplaya Yungas Bolivia LA2811 Cerro Huayrapampa Apurimac Peru LA2814 Ccascani, Sandia Puno Peru LA2841 Chinuna Amazonas Peru LA2842 Santa Rita San Martin Peru LA2843 mercado San Martin Peru LA2844 Shanhao San Martin Peru LA2845 Mercado Moyobamba San Martin Peru LA2871 Chamaca La Paz Bolivia LA2873 Lote Pablo Luna #2 La Paz Bolivia LA2874 Playa Ancha La Paz Bolivia LA2933 Jipijapa Manabi Ecuador LA2977 Belen Beni Bolivia LA2978 Belen Beni Bolivia LA3135 Pinal del Jigue Holguin LA3136 Arroyo Rico Holguin Cuba LA3137 Pinares de Mayari Holguin Cuba LA3138 El Quemada Holguin Cuba LA3139 San Pedro de Cananova Holguin Cuba LA3140 Los Platanos Holguin Cuba LA3141 Guira de Melena La Habana Cuba LA3162 N of Copan Honduras LA3452 CATIE, Turrialba Turrialba Costa Rica LA3633 Botanical garden Ghana LA3842 El Limon, Maracay Aragua Venezuela

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Solanum lycopersicum (L. esculentum var. cerasiforme) LA3843 El Limon, Maracay Aragua Venezuela LA3844 Algarrobito Guarico Venezuela LA4352 Bamoa Sinaloa Mexico LA4353 Guasave Sinaloa Mexico

Solanum neorickii (L. parviflorum) LA0247 Chavinillo Huanuco Peru LA0735 Huariaca Huanuco Peru LA1319 Apurimac Peru LA1321 Curahuasi Apurimac Peru LA1322 Limatambo Cusco Peru LA1326 Rio Pachachaca Apurimac Peru LA1329 Yaca Apurimac Peru LA1626A Mouth of Rio Rupac Ancash Peru LA1716 Huancabamba Piura Peru LA2072 Huanuco Huanuco Peru LA2073 Huanuco, N of San Rafael Huanuco Peru LA2074 Huanuco Huanuco Peru LA2075 Huanuco Huanuco Peru LA2113 La Toma Loja Ecuador LA2133 Ona Azuay Ecuador LA2190 Tialango Amazonas Peru LA2191 Campamento Ingenio Amazonas Peru LA2192 Pedro Ruiz Amazonas Peru LA2193 Churuja Amazonas Peru LA2194 Chachapoyas West Amazonas Peru LA2195 Caclic Amazonas Peru LA2197 Caclic ‐ Luya Amazonas Peru LA2198 Chachapoyas East Amazonas Peru LA2200 Choipiaco Amazonas Peru LA2201 Pipus Amazonas Peru LA2202 Tingobamba Amazonas Peru LA2315 Sargento Amazonas Peru LA2317 Zuta Amazonas Peru LA2318 Lima Tambo Amazonas Peru LA2319 Chirico Amazonas Peru LA2325 Above Balsas Amazonas Peru LA2403 Wandobamba Huanuco Peru LA2613 Matichico‐San Rafael Huanuco Peru LA2614 San Rafael Huanuco Peru LA2615 Ayancocho Huanuco Peru LA2639A Puente Cunyac Apurimac Peru LA2641 Nacchera Apurimac Peru LA2727 Ona Azuay Ecuador LA2847 Suyubamba Amazonas Peru

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Solanum neorickii (L. parviflorum) LA2848 Pedro Ruiz Amazonas Peru LA2862 Saraguro‐Cuenca Azuay Ecuador LA2865 Rio Leon Azuay Ecuador LA2913 Uchucyaco ‐ Hujainillo Huanuco Peru LA4020 Gonozabal Loja Ecuador LA4021 Guancarcucho Azuay Ecuador LA4022 Pueblo Nuevo Azuay Ecuador LA4023 Paute Azuay Ecuador

Solanum ochranthum LA2118 San Lucas Loja Ecuador LA2160 Acunac Cajamarca Peru LA2161 Cruz Roja Cajamarca Peru LA2162 Yatun Cajamarca Peru LA2166 Pacopampa Cajamarca Peru LA2203 Pomacochas Amazonas Peru LA2682 Chinchaypujio Cusco Peru

Solanum pennellii (L. pennellii or L. pennellii var. puberulum) LA0716 Atico Arequipa Peru LA0750 Palpa to Nazca Ica Peru LA0751 Sisacaya Lima Peru LA1272 Pisaquera Lima Peru LA1273 Cayan Lima Peru LA1275 Quilca road junction Lima Peru LA1277 Trapiche Lima Peru LA1282 Sisacaya Lima Peru LA1297 Pucara Lima Peru LA1299 Santa Rosa de Quives Lima Peru LA1302 Quita Sol Ica Peru LA1303 Pampano Huancavelica Peru LA1340 Pacaran Lima Peru LA1356 Moro Ancash Peru LA1367 Santa Eulalia Lima Peru LA1376 Sayan Lima Peru LA1515 Sayan to Churin Lima Peru LA1522 Quintay Lima Peru LA1523* Irrigacion Santa Rosa Lima Peru LA1649 Molina Ica Peru LA1656 Marca to Chincha Ica Peru LA1657 Buena Vista to Yautan Ancash Peru LA1674 Toparilla Canyon Lima Peru LA1693 Quebrada Machuranga Lima Peru LA1724 La Quinga Ica Peru LA1732 Rio San Juan Huancavelica Peru

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Solanum pennellii (L. pennellii or L. pennellii var. puberulum) LA1733 Rio Canete Lima Peru LA1734 Rio Canete Lima Peru LA1735 Rio Canete Lima Peru LA1773 Rio Casma Ancash Peru LA1809 El Horador (playa) Piura Peru LA1911 Locari Ica Peru LA1912 Cerro Locari Ica Peru LA1920 Cachiruma Ayacucho Peru LA1926 Agua Perdida Ica Peru LA1940 Rio Atico, Km 26 Arequipa Peru LA1941 Rio Atico, Km 41 Arequipa Peru LA1942 Rio Atico, Km 54 Arequipa Peru LA1943 Rio Atico, Km 61 Arequipa Peru LA1946 Caraveli Arequipa Peru LA2560 Santa to Huaraz Ancash Peru LA2580 Valle de Casma Ancash Peru LA2657 Bayovar Piura Peru LA2720* Pacaran‐Catahuasi Lima Peru LA2963 Acoy Arequipa Peru

Solanum peruvianum (L. peruvianum) LA0098 Chilca Lima Peru LA0111 Supe Lima Peru LA0153 Culebras Ancash Peru LA0370 Hacienda Huampani Lima Peru LA0371 Supe Lima Peru LA0372 Culebras #1 Ancash Peru LA0374 Culebras #2 Ancash Peru LA0445 Chincha #2 Ica Peru LA0446 Lomas de Atiquipa Arequipa Peru LA0448 Chala Arequipa Peru LA0453 Yura #2 Arequipa Peru LA0454 Tambo Arequipa Peru LA0455 Tambo Arequipa Peru LA0462 Sobraya Arica‐Parinacota Chile LA0464 Hacienda Rosario Arica‐Parinacota Chile LA0752 Sisicaya Lima Peru LA1161 Huachipa Lima Peru LA1270 Pisiquillo Lima Peru LA1278 Trapiche Lima Peru LA1333 Loma Camana Arequipa Peru LA1336 Atico Arequipa Peru LA1337 Atiquipa Arequipa Peru LA1368 San Jose de Palla Lima Peru LA1474 Lomas de Camana Arequipa Peru

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Solanum peruvianum (L. peruvianum) LA1475 Fundo 'Los Anitos', Barranca Lima Peru LA1513 Atiquipa Arequipa Peru LA1517 Irrigacion Santa Rosa Lima Peru LA1537 Azapa Valley Arica‐Parinacota Chile LA1556 Hacienda Higuereta Lima Peru LA1616 La Rinconada Lima Peru LA1675 Toparilla Canyon Lima Peru LA1692 Putinza Lima Peru LA1913 Tinguiayog Ica Peru LA1929 La Yapana Ica Peru LA1935 Lomas de Atiquipa Arequipa Peru LA1947 Puerto Atico Arequipa Peru LA1949 Las Calaveritas Arequipa Peru LA1951 Ocona Arequipa Peru LA1952* Lomas de Camana Arequipa Peru LA1954 Arequipa Peru LA1955 Matarani Arequipa Peru LA1975 Desvio Santo Domingo Lima Peru LA1977 Orcocoto Lima Peru LA1989 (self‐compatible selection) LA2573 Valle de Casma Ancash Peru LA2581 Chacarilla (autotetraploid) Arica‐Parinacota Chile LA2732 Moquella Tarapaca Chile LA2742 Camarones‐Guancarane Arica‐Parinacota Chile LA2744 Sobraya Arica‐Parinacota Chile LA2745 Pan de Azucar Arica‐Parinacota Chile LA2770 Lluta valley Arica‐Parinacota Chile LA2834 Hacienda Asiento Ica Peru LA2955B Quistagama‐Quisama Tarapaca Chile LA2957B Pozo Tarapaca Chile LA2958* Caleta Vitor Arica‐Parinacota Chile LA2959 Chaca to Caleta Vitor Arica‐Parinacota Chile LA2964 Quebrada de Burros Tacna Peru LA3218 Quebrada Guerrero Arequipa Peru LA3220 Cocachacra Arequipa Peru LA3640 Mexico LA3858 Canta Lima Peru LA3900 (CMV tolerant selection) LA4125 Camina Tarapaca Chile LA4317 Rio Lluta, desembocadura Arica‐Parinacota Chile LA4318 Sora ‐ Molinos, Rio Lluta Arica‐Parinacota Chile LA4325 Caleta Vitor Arica‐Parinacota Chile LA4328 Pachica, Rio Camarones Arica‐Parinacota Chile LA4445 Azapa Valley, 27 km from Arica Arica‐Parinacota Chile LA4446 Azapa Valley, Km 37 from Arica Arica‐Parinacota Chile

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Solanum peruvianum (L. peruvianum) LA4447 Azapa Valley, Km 27 and Km 37 from Tarapaca Chile

Solanum pimpinellifolium (L. pimpinellifolium) LA0100 La Cantuta (Rimac Valley) Lima Peru LA0114 Pacasmayo La Libertad Peru LA0121 Trujillo La Libertad Peru LA0122 Poroto La Libertad Peru LA0369 La Cantuta (Rimac Valley) Lima Peru LA0373 Culebras #1 Ancash Peru LA0375 Culebras #2 Ancash Peru LA0376 Hacienda Chiclin La Libertad Peru LA0381 Pongo La Libertad Peru LA0391 Magdalena (Rio Jequetepeque) Cajamarca Peru LA0397 Hacienda Tuman Lambayeque Peru LA0398 Hacienda Carrizal Cajamarca Peru LA0400 Hacienda Buenos Aires Piura Peru LA0411 Pichilingue Los Rios Ecuador LA0412 Pichilingue Los Rios Ecuador LA0413 Cerecita Guayas Ecuador LA0417 Punta Polvora, Isla Puna Guayas Ecuador LA0418 Daule Guayas Ecuador LA0420 El Empalme Guayas Ecuador LA0442 Sechin Ancash Peru LA0443 Pichilingue Los Rios Ecuador LA0480 Hacienda Santa Inez Ica Peru LA0722 Trujillo La Libertad Peru LA0753 Lurin Lima Peru LA1236 Hotel Tinalandia, Santo Domingo Pichincha Ecuador LA1237 Atacames Esmeraldas Ecuador LA1242 Los Sapos Guayas Ecuador LA1243 Murillo finca Guayas Ecuador LA1245 Santa Rosa El Oro Ecuador LA1246 La Toma Loja Ecuador LA1248 Hacienda Monterrey Loja Ecuador LA1256 Naranjal Guayas Ecuador LA1257 Las Mercedes Guayas Ecuador LA1258 Voluntario de Dios Guayas Ecuador LA1259 Catarama Los Rios Ecuador LA1260 Pueblo Viejo Los Rios Ecuador LA1261 Babahoyo Los Rios Ecuador LA1262 Milagro Empalme Guayas Ecuador LA1263 Barranco Chico Guayas Ecuador LA1269 Pisiquillo Lima Peru LA1279 Cieneguilla Lima Peru LA1280 Chontay Lima Peru

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Solanum pimpinellifolium (L. pimpinellifolium) LA1301 Hacienda San Ignacio Ica Peru LA1332 Nazca area? Ica Peru LA1335 Pescadores Arequipa Peru LA1341 Huampani Lima Peru LA1342 Casma Ancash Peru LA1343 Puente Chao La Libertad Peru LA1344 Laredo La Libertad Peru LA1345 Samne La Libertad Peru LA1348 Pacasmayo La Libertad Peru LA1349 Cuculi Lambayeque Peru LA1355 Nepena Ancash Peru LA1357 Jimbe Ancash Peru LA1359 La Crau Ancash Peru LA1370 San Jose de Palla Lima Peru LA1371 Santa Eulalia Lima Peru LA1374 El Ingenio Ica Peru LA1375 San Vicente de Canete Lima Peru LA1380 Chanchape Piura Peru LA1381 Naupe Lambayeque Peru LA1382 Chachapoyas to Balsas Amazonas Peru LA1383 Chachapoyas to Bagua Amazonas Peru LA1384 Quebrada Parca Lima Peru LA1416 Las Delicias Pichincha Ecuador LA1428 La Estancilla Manabi Ecuador LA1466 Chongoyape Lambayeque Peru LA1469 El Pilar, Olmos Lambayeque Peru LA1470 Motupe to Desvio Olmos‐Bagua Lambayeque Peru LA1471 Motupe to Jayanca Lambayeque Peru LA1472 Quebrada Topara Lima Peru LA1478 Santo Tome (Pabur) Piura Peru LA1514 Sayan to Churin Lima Peru LA1520 Sayan to Churin Lima Peru LA1521 El Pinon, Asia Lima Peru LA1547 Chota to El Angel Carchi Ecuador LA1561 San Eusebio Lima Peru LA1562 Cieneguilla Lima Peru LA1571 San Jose de Palle Lima Peru LA1572 Hacienda Huampani Lima Peru LA1573 Nana Lima Peru LA1575 Huaycan Lima Peru LA1576 Manchay Alta Lima Peru LA1577 Cartavio La Libertad Peru LA1578 Santa Marta La Libertad Peru LA1579 Colegio Punto Cuatro #1 Lambayeque Peru LA1580 Colegio Punto Cuatro #2 Lambayeque Peru

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Solanum pimpinellifolium (L. pimpinellifolium) LA1581 Punto Cuatro Lambayeque Peru LA1582 Motupe Lambayeque Peru LA1583 Tierra de la Vieja Lambayeque Peru LA1584 Jayanca to La Vina Lambayeque Peru LA1585 Cuculi Lambayeque Peru LA1586 Zana, San Nicolas La Libertad Peru LA1587 San Pedro de Lloc La Libertad Peru LA1588 Laredo to Barraza La Libertad Peru LA1589 Viru ‐ Galunga La Libertad Peru LA1590 Viru to Tomaval La Libertad Peru LA1591 Ascope La Libertad Peru LA1592 Moche La Libertad Peru LA1593 Puente de Chao La Libertad Peru LA1594 Cerro Sechin Ancash Peru LA1595 Nepena to Samanco Ancash Peru LA1596 Santa to La Rinconada Ancash Peru LA1597 Rio Casma Ancash Peru LA1598 Culebras to La Victoria Ancash Peru LA1599 Huarmey Ancash Peru LA1600 Las Zorras, Huarmey Ancash Peru LA1601 La Providencia Lima Peru LA1602 Punchauca Lima Peru LA1603 Quilca Lima Peru LA1604 Horcon Lima Peru LA1605 Canete ‐ San Antonio Lima Peru LA1606 Tambo de Mora Ica Peru LA1607 Canete ‐ La Victoria Lima Peru LA1608 Canete ‐ San Luis Lima Peru LA1610 Asia ‐ El Pinon Lima Peru LA1611 Rio Mala Lima Peru LA1612 Rio Chilca Lima Peru LA1613 Hacienda Santa Eusebia Lima Peru LA1614 Huaura ‐‐ Pampa Chumbes Lima Peru LA1615 Piura to Simbala Piura Peru LA1617 Tumbes South Tumbes Peru LA1618 Tumbes North Tumbes Peru LA1628 Huanchaco La Libertad Peru LA1629 Barrancos de Miraflores Lima Peru LA1630 Fundo La Palma Ica Peru LA1631 Planta Envasadora San Fernando La Libertad Peru LA1633 Coop. Ica Peru LA1634 Fundo Bogotalla #1 Ica Peru LA1635 Fundo Bogotalla #2 Ica Peru LA1636 Laran Ica Peru LA1637 La Calera Ica Peru

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Solanum pimpinellifolium (L. pimpinellifolium) LA1638 Fundo El Portillo Lima Peru LA1645 Banos de Miraflores Lima Peru LA1651 Vivero, La Molina Lima Peru LA1652 Cieneguilla Lima Peru LA1659 Pariacoto Ancash Peru LA1660 Yautan to Pariacoto Ancash Peru LA1661 Esquina de Asia Lima Peru LA1670 Rio Sama Tacna Peru LA1676 Fundo Huadquina, Topara Ica Peru LA1678 San Juan Lucumo de Topara Ica Peru LA1679 Tambo de Mora Ica Peru LA1680 La Encanada Lima Peru LA1682 Montalban ‐ San Vicente Lima Peru LA1683 Miramar Piura Peru LA1684 Chulucanas Piura Peru LA1685 Marcavelica Piura Peru LA1686 Valle Hermoso #1 Piura Peru LA1687 Valle Hermoso #2 Piura Peru LA1688 Pedregal Piura Peru LA1689 Castilla #1 Piura Peru LA1690 Castilla #2 Piura Peru LA1697 Hacienda Quiroz, Santa Anita Lima Peru LA1719 E of Arenillas El Oro Ecuador LA1720 Yautan Ancash Peru LA1728 Rio San Juan Ica Peru LA1729 Rio San Juan Ica Peru LA1742 Olmos‐Marquina Lambayeque Peru LA1781 Bahia de Caraquez Manabi Ecuador LA1921 Majarena Ica Peru LA1923 Coop. Ica Peru LA1924 Piedras Gordas Ica Peru LA1925 Pangavari, Nazca Ica Peru LA1933 Jaqui Arequipa Peru LA1936 Huancalpa Arequipa Peru LA1950 Pescadores Arequipa Peru LA1987 Viru ‐ Fundo Luis Enrique La Libertad Peru LA1992 Pichicato Lima Peru LA1993 Chicama Valley? Lima Peru LA2093 La Union El Oro Ecuador LA2096 Playa, near Catacocha Loja Ecuador LA2097 Macara Loja Ecuador LA2102 El Lucero Loja Ecuador LA2112 Hacienda Monterrey Loja Ecuador LA2145 Juan Montalvo Los Rios Ecuador LA2146 Hacienda Limoncarro La Libertad Peru

Page 57 of 64 STOCK LISTS TGC REPORT VOLUME 63, 2013

Solanum pimpinellifolium (L. pimpinellifolium) LA2147 Yube Cajamarca Peru LA2149 Puente Muyuna Cajamarca Peru LA2170 Pai Pai Cajamarca Peru LA2173 Cruz de Huayquillo Cajamarca Peru LA2176 Timbaruca Cajamarca Peru LA2178 Tororume Cajamarca Peru LA2179 Tamboripa ‐ La Manga Cajamarca Peru LA2180 La Coipa Cajamarca Peru LA2181 Balsa Huaico Cajamarca Peru LA2182 Cumba Amazonas Peru LA2183 Corral Quemado Amazonas Peru LA2184 Bagua Amazonas Peru LA2186 El Salao Amazonas Peru LA2187 La Caldera Amazonas Peru LA2188 Manchungal #1 Amazonas Peru LA2189 Manchungal #2 Amazonas Peru LA2335 (autotetraploid) LA2340 (autotetraploid) LA2345 (autodiploid) LA2346 (autodiploid) LA2347 (autodiploid) LA2348 Trujillo La Libertad Peru LA2389 Tembladera Cajamarca Peru LA2390 Chungal Cajamarca Peru LA2391 Chungal to Monte Grande Cajamarca Peru LA2401 Moxeque Ancash Peru LA2412 Fundo Don Javier, Chilca Lima Peru LA2533 Lomas de Latillo Lima Peru LA2534* Vista Florida, Lambayeque Peru LA2535* Patapo Lambayeque Peru LA2536* Patapo‐La Cria Lambayeque Peru LA2537* Malpaso Lambayeque Peru LA2538* Malpaso Lambayeque Peru LA2539* Cuculi Lambayeque Peru LA2540* La Cria Lambayeque Peru LA2543* Pacanguilla‐Pacasmayo La Libertad Peru LA2544* Pacasmayo‐Huabal La Libertad Peru LA2545* Tembladera Cajamarca Peru LA2546* Tonan Cajamarca Peru LA2547* Yatahual Cajamarca Peru LA2549* La Muyuna Cajamarca Peru LA2576 Valle de Casma Ancash Peru LA2578 Tuturo Ancash Peru LA2585 (autotetraploid) LA2628 Echarate/Tornillochayoc Cusco Peru

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Solanum pimpinellifolium (L. pimpinellifolium) LA2645 Desvio Chulucanas Piura Peru LA2646 Chalaco Piura Peru LA2647 Morropon‐Chalaco Piura Peru LA2649* Chulucanas Piura Peru LA2652 Piura Peru LA2653 San Francisco de Chocon Querecotillo Piura Peru LA2655 La Huaca ‐ Sullana Piura Peru LA2656 Suarez Tumbes Peru LA2659 Castilla, Univ. Nac. de Piura Piura Peru LA2718 Chilca Lima Peru LA2725 Ica Peru LA2805 (‘Indehiscent currant’) LA2831 Rio Nazca Ica Peru LA2832 Chichictara Ica Peru LA2833 Hacienda Asiento Ica Peru LA2836 Rio Aja Ica Peru LA2839 Tialango Amazonas Peru LA2840 San Hilarion de Tomaque Amazonas Peru LA2850 Santa Rosa, Manta Manabi Ecuador LA2851 La Carcel de Montecristo Manabi Ecuador LA2852 Cirsto Rey de Charapoto Manabi Ecuador LA2853 Experiment Station, Portoviejo‐INIAP Manabi Ecuador LA2854 Jipijapa Manabi Ecuador LA2857 Isabela: Puerto Villamil Galapagos Ecuador LA2866 Via a Amaluza Loja Ecuador LA2914A Urb. La Castellana, Surco Lima Peru LA2914B Urb. La Castellana, Surco Lima Peru LA2915 El Remanso de Olmos Lambayeque Peru LA2934 Carabayllo Lima Peru LA2966 La Molina Lima Peru LA2974 La Libertad Peru LA2982 Chilca #1 Lima Peru LA2983 Chilca #2 Lima Peru LA3123 Santa Cruz: summit Galapagos Ecuador LA3158 Los Mochis Sinaloa Mexico LA3159 Los Mochis Sinaloa Mexico LA3160 Los Mochis Sinaloa Mexico LA3161 Los Mochis Sinaloa Mexico LA3859 (TYLCV resistant selection) LA4138 El Corregidor, La Molina Lima Peru LA4431 (sun in LA1589)

Solanum sitiens (S. rickii) LA1974 Chuquicamata Antofagasta Chile LA2876 Chuquicamata Antofagasta Chile

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Solanum sitiens (S. rickii) LA2877 El Crucero Antofagasta Chile LA2878 Mina La Exotica Antofagasta Chile LA2885 Caracoles Antofagasta Chile LA4105 Mina La Escondida Antofagasta Chile LA4110 Mina San Juan Antofagasta Chile LA4112 Aguada Limon Verde Antofagasta Chile LA4113 Estacion Cere Antofagasta Chile LA4114 Pampa Carbonatera Antofagasta Chile LA4115 Quebrada desde Cerro Oeste de Paqui Antofagasta Chile LA4116 Quebrada de Paqui Antofagasta Chile LA4331 Cerro Quimal Antofagasta Chile

*Previously inactive, newly regenerated accession

Page 60 of 64 MEMBERSHIP LIST TGC REPORT VOLUME 63, 2013

Membership List

Aarden, Harriette Monsanto Holland BV, Dept. Tomato Breeding, LAAN VAN BRAAT, 21, DELFT, THE NETHERLANDS 2627CL [email protected] Ballardini, Massimiliano Edasem Spa, Via G. Marconi 56, Casaleone, Verona, ITALY, 37052 [email protected] Barker, Susan University of Western Australia, School of Plant Biology M084, 35 Stirling Hwy, Crawley, WA, AUSTRALIA, 6009 [email protected] Brown, Loretta Rijk Zwaan USA, 19040 Portola Dr Ste B, Salinas, CA, USA, 93908 [email protected] Buonfiglioli, Carlo Della Rimembranze nr. 6A, San Lazzaro di Savena, Bologna, ITALY 40068 [email protected] Chen, Dei Wei Bucolic Seeds Co. Ltd., P.O. 2-39, Tantzu, Taichung Co., TAIWAN, 427 [email protected] Chetelat, Roger University of California, Dept of Veg Crops, One Shields Ave, Davis, CA, USA 95616-8746 [email protected] deHoop, Simon Jan Hortigenetics Research LTD S.E. Asia, No. 7 Moo 9 Maefackmai, Sansai district, Chiang Mai, THAILAND 50290 [email protected] Galvez, Hayde U Philippines Los Banos, Genetics Lab, CSC-1PB, CA, UP Los Banos College Laguna, PHILIPPINES 4031 [email protected] Haga, Emily Johnny's Selected Seeds, 184 Foss Hill Rd, Albion, ME, USA, 04910 [email protected] Hanson, Peter AVRDC , PO Box 42, Shanhua, Tainan, TAIWAN, REPUBLIC of 741 [email protected] Herlaar, Fritz Enza Zaden Export B.V., Research and Development, P.O. Box 7, Enkhuizen, THE NETHERLANDS, 1600 AA [email protected] Holloway, Heather M. Nightshade LLC, 816 E. Pennsylvania Dr., Boise, ID, USA, 83706 [email protected] Hoogstraten, Jaap Seminis Veg Seeds, Postbus 97, 6700 AB Wageningen, THE NETHERLANDS [email protected] Hutton, Sam University of Florida, Gulf Coast Research and Education Center, 14625 County Rd 672, Wimuama, FL, USA, 33598 [email protected] Inai, Shuji Nippon Del Monte Corp., Research and Development, 3748 Shimizu-Cho, Numata-shi, Gunma-ken, JAPAN, 378-0016 [email protected] Joly, Marie-Pierre Vilmorin, Route du Manoir, La Menitre, , 49250 [email protected] Kaplan, Boaz Nirit Seeds LTD Ltd, P.O. Box 95, Moshav Hadar, Am, ISRAEL, 42935 [email protected] Karwa, Anup Krishidhan Veg Seeds Private LTD, Krishidhan Bhanwan,, Additional MIDC Jalna, Maharashtra, INDIA, 431203 [email protected] Kobori, Romulo F. Sakata Seed Sudamerica Ltda, PO Box 427, Av Dr. Plinio Salgado N, 4320, Bairro, Braganca Paulista, Sao Paulo, BRAZIL, 12-906-840 [email protected]

Page 61 of 64 MEMBERSHIP LIST TGC REPORT VOLUME 63, 2013

Kuehn, Michael HM Clause, 25757 CR 21A, Esparto, CA, USA, 95627 [email protected] Massoudi, Mark Ag Biotech Inc., P.O. Box 1325, San Juan Bautista, CA, USA, 95045 [email protected] Maxwell, Douglas P. Max- Tach Services, 7711 Midtown Rd, Verona, WI, USA, 53593 [email protected] Moisson, Chloe Technisem, Zone d' activite Anjou Actiparc de Jumelles, Longue Jumelles, FRANCE, 49160 [email protected] Myers, Jim Oregon State University, Dept. of Horticulture, rm 4017, Ag & Life Sci Bldg., Corvallis, OR, USA, 97331 [email protected] Nakamura, Kosuke Kagome Co. Ltd., 17 Nishitomiyama, Nasushiobarashi, Tochigi, JAPAN, 329-2762 [email protected] Nukal, Balaji SeedWorks, 155 Ocean Lane Dr, #515, Key Biscayne, FL 33149 [email protected] O'Brochta, William Roanoke Valley Governor's School, 3226 Pearwood Dr, Roanoke, VA, USA, 24014 [email protected] Peters, Susan Nunhems USA, 7087 E. Peltier Rd., Acampo, CA, USA, 95220 [email protected] Randhawa, Parm California Seed and Plant Lab , 7877 Pleasant Grove Rd, Elverta, CA, USA, 95626 [email protected] Rascle, Christine CLAUSE, Centre de Recherche, Domaine de Maninet, Route de Beaumont, Valence, FRANCE, 26000 [email protected] Schuit, C.A. Bejo Zaden B.V., Molecular Market Technology & Genomics, Trambaan 2, Warmenhuizen, CZ, Netherlands, 1747 [email protected] Scott, Jay University of Florida, Tomato Breeders, 14625 County Rd 672, Wimauma, Fl, USA, 33598 [email protected] Serquen, Felix Syngenta Seeds, 21435 Road 98, Woodland, CA, USA, 95695 [email protected] Sharma, Sundrish Syngenta Seeds, Inc., 10290 Greenway Rd, Naples, FL, USA, 34114 [email protected] Shekaste band, Reza University of Florida, Gulf Coast Research and Education Center, 14625 County Rd 672 Wimuama FL USA 33598 [email protected] Shintaku, Yurie 2-10-2, Shimizu, Suginami-ku, Tokyo, JAPAN , 167-0033 [email protected] Stack , Stephen Colorado State U, Dept of Biology, 1878 Campus Delivery, Fort Collins, CO, USA, 80523-1878 [email protected] Stommel, Ph.D., John USDA-ARS, Genetic Improvement Fruits & Vegetables Laboratory, Bldg. 010A, BARC-West, 10300 Baltimore Ave., Beltsville, MD, USA, 20705 [email protected] Thome, Catherine United Genetics Seeds Co., 764 Carr Ave., Aromas, CA, USA, 65004 [email protected] Tikoo, Surinder K. Tierra Seed Science Pvt. Ltd., Breeding & Develoopment, Malaxmi Courtyard, Khajaguda, Golconda Post, Hyderabad, INDIA, 500 008 [email protected]

Page 62 of 64 MEMBERSHIP LIST TGC REPORT VOLUME 63, 2013 van der Knaap, Koen Axia Vegetable Seeds, Delft, The Netherlands, 262gHH [email protected] van der Knaap , Ben FutureSupport Consultancy [email protected] van Vuuren, Ansa Starke Ayres Seed (Pty) Ltd, Pepper and Tomato Researcher, P.O. Box 14366, Gauteng, BREDELL, SOUTH AFRICA , 1625 [email protected] Vecchio, Franco Nunhems Italy SRL, Via Ghiarone 2, Sant' Agata, Bolognese (BO), ITALY, 40019 Verschave, Philippe Vilmorin, Route de Meynes, Ledenon, FRANCE, 30210 [email protected] Volpin, Hanne Hazera Genetics Ltd., R&D Division, M.P. Lachish Darom, Lachish Darom, M.P., ISRAEL, 79354 [email protected] Williamson, Valerie UC Davis, Dept. of Nematology, 1 Shields Ave, Davis CA, USA, 95616 [email protected] Zhiqi Jia Henan Agricultural University room 313, no 2 building, No 95, Wenhua Rd, Jinshui District, Zhengzhou City, Henan Province, CHINA, 450002 [email protected]

Page 63 of 64 AUTHOR INDEX TGC REPORT VOLUME 63, 2013

AUTHOR INDEX

Anderson, Lorinda K...... 5 Chetelat, R. T ...... 31 Chung, Chi-Yun ...... 22 de Jong, Hans...... 5 Giovannoni, James J...... 5 Goicoechea, José Luis ...... 5 Hanson, Peter ...... 15 Ho, Fang-I ...... 15, 22 Hua, Axin ...... 5 Hutton, S.F………………………………………………………………………32 Ledesma, Dolores ...... 15 Lu, Shu-Fen ...... 15 Roe, Bruce A...... 5 Scott, J.W…………………………………………………………………..31, 32 Shearer, Lindsay A...... 5 Smit, Sandra ...... 5 Stack, Stephen M ...... 5 Tan, Chee-Wee ...... 15 Wang, Jaw-Fen ...... 15, 22

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